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CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY
II
TAXONOMY AND
EOGRAPHICAL DISTRIBUTION
OF THE GENUS MILESIA
BY
JOSEPH HORACE FAULL
WITH NINE PLATES
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PUBLISHED BY
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JAMAICA PLAIN, MASS., U.S.A.
1932
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HARVARD UNIVERSITY
THE GIFT OF
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY
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No. I. The Hypodermataceae of Conifers. By G. D.
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No. II. Taxonomy and Geographical Distribution of
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TAXONOMY AND GEOGRAPHICAL
DISTRIBUTION OF THE
GENUS MILESIA
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY
II
TAXONOMY AND
GEOGRAPHICAL DISTRIBUTION
OF THE GENUS MILESIA
BY
JOSEPH HORACE FAULL
WITH NINE PLATES
PUBLISHED BY
THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY
JAMAICA PLAIN, MASS., U.S.A.
1932
Contributions from the Arnold Arboretum
of Harvard University
No. 2, pp. 1-138, plates i-ix
Issued October 1, 1932
TABLE OF CONTENTS
INTRODUCTION . 5
NOMENCLATURE . 5
HISTORICAL RLSUML . 11
HOSTS . 16
GEOGRAPHICAL DISTRIBUTION . 18
THE TAXONOMY OF MILESIA . 21
The Genus Milesia . 22
Key to the Species of Milesia . 24
I. Species on Asplenium . 27
II. Species on Blechnum . 37
Til Species on Cryptogramma . 46
IV. Species on Dennstaedtia . 48
V. Species on Dryopteris . 49
VI. Species on Histtopteris . 73
VII. Species on Nephrolepis . 74
VIII. Species on Pellaea . 77
IX. Species on Polypodium . 81
X. Species on Polystichum . 100
XI. Species on Scolopendrium . 113
FERN HOSTS AND THE SPECIES OF MILESIA THAT OCCUR ON
THEM . 119
THE SPECIES OF MILESIA, THE TWO GENERATIONS OF
WHICH ARE KNOWN, AND THEIR ALTERNATE HOSTS . 120
SPECIES OF MILESIA AND THEIR FERN HOSTS . 120
EXCLUDED SPECIES . 121
ACKNOWLEDGMENTS . 123
SUMMARY . 124
BIBLIOGRAPHY . 126
ADDENDUM TO “V. Species on Dryopteris” . 129
EXPLANATION OF PLATES . 131
INDEX
135
)
TAXONOMY AND GEOGRAPHICAL
DISTRIBUTION OF THE
GENUS MILESIA
INTRODUCTION
The Family Melampsoraceae as at present constituted is an
exceedingly fascinating group of rusts because of the great
variety of forms of which it is composed and because of the
many intricate phylogenetic problems it offers. Of its Sub¬
families, the Pucciniastreae is accorded by many uredinologists
a foremost position in these respects. Standing out among the
Pucciniastreae as of peculiar interest are the so-called “Fern
rusts,’7 the Uredinopsatae of Arthur, of which there are three
recognized genera, namely, Uredinopsis, Milesia and Hyalop-
sora. The uredinial and the telial stages of all species of these
genera are restricted to ferns, and the aecial, of all whose life
histories are known, to Firs (Abies). Uredinopsis is thought
by many uredinologists to represent a primitive genus of rusts,
and Milesia is apparently a close relative. Both are unique in
that their constituents show not a trace of pigmentation; they
are colorless throughout, and their spores are white in mass.
This paper, the second of a series on the Pucciniastreae (16),
is devoted to the taxonomy and the geographical distribution
of the genus Milesia.
NOMENCLATURE
The genus Milesia was established by White (48) in 1877,
and was founded on the uredinial stage of an hitherto unde-
scribed species of rust discovered at various stations through¬
out Perthshire, Scotland, on fronds of Poly podium vulgar e.
It was named in honor of Miles Joseph Berkeley. The species
was designated “ Milesia polypodii B. White sp. n.” White’s
definition of both the genus and the species applied to the ure¬
dinial phase only. This was the only spore phase alluded to,
and without saying which one he thought it was, he ventured
to remark that it was Endophyllum-like. But his descriptions
6
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
were clear, accurate, and correctly illustrated by drawings.
No doubt can exist as to the identity of his materials. He
commented on the smallness of the uredinia, keenly observed
the thin and very inconspicuous peridium, noted that the spores
were copiously discharged through a minute ostiolum, de¬
scribed the spores as distinctly spiny, gave their true lineal
measurements, and did not fail to mention that they were
white. Knowing nothing of nuclear constitution nor of the
teliospores, the conclusion that his fungus was a rust resulted,
of course, from inference and not proof. It was, however, a
noteworthy inference because with few exceptions, such for
example as a similar conclusion reached two years previously
by Oudemans (39) with respect to a similar rust on Scolopen-
drium vulgar e, white-spored fungi on ferns had been inter¬
preted as species of Gloeophyllum , Septoria , etc., and not as
rusts. His collections, which he stated showed abundant spore
production, were made in June and July on “living but fading”
leaves, that is on overwintered fronds. We now know that
this habit of copious production of uredospores on overwin¬
tered fronds is characteristic of the species of Milesia in gen¬
eral, as contrasted with those of Uredinopsis, the only white-
spored rusts with which they might be confused in the ure-
dinial phase, quite apart from the ordinary differences in ure-
dospore shapes and markings.
The very same rust was redescribed from Germany by Sy-
dow (42) twenty-six years later (1903) under the name Me-
lampsorella Dietelima Syd., nov. spec. The only important
addition to White’s earlier description was a statement for the
first time concerning teliospores. Sydow gave a detailed de¬
scription of the uredinia and uredospores, and stated that he
had observed the rare occurrence of many-celled, hyaline,
intracellular teliospores. He made no allusion in any way to
Milesia Polypodii White or to White’s earlier description of
the rust. The next significant mention of this rust on Poly¬
podium vulgar e was made by Arthur (1) two years later in a
paper presented before the International Botanical Congress
at Vienna in 1905; Arthur cited it as the type of the genus
Milesia, employing the earlier name, Milesia Polypodii White,
and listed several species of Uredinopsis as members likewise
of the genus Milesia. Four years later still Magnus (37), in
1932]
FAULL, THE GENUS MILESIA
7
the course of his studies on certain genera of the Melamp-
soraceae re-christened the rust “ Milesina Dieteliana ,” remov¬
ing it from Melampsorella, but retaining the specific name
“Dieteliana” in preference to accepting White’s “Polypodii.”
This disposition of the species by Magnus has continued to
receive wide though not universal acceptance (as opposed see
Arthur 5, p. 685, and Clements and Shear in “The Genera of
Fungi,” 1931, p. 338, who retain White’s Milesia Polypodii”)
on the ground that the specific name “Dieteliana” published
in 1903, is based on the teliosporic, or “perfect” stage, and so
takes priority over the corresponding name “Polypodii” of
White according to the International Rules of Nomenclature
formulated by the Brussels International Botanical Congress
of 1910.
Coincident with the acceptance of the specific name “Diete¬
liana” Magnus rejected White’s genus Milesia, on the assump¬
tion that it was a monotypic genus that included a single
species, founded on an “imperfect” spore stage, the generic
identity of which was indeterminable. He made no reference
to Arthur’s citation of 1905. Magnus included in Milesina
two species, transferred from Melampsorella as then consti¬
tuted, namely, M. Kriegeriana and M. Feurichii, and in a foot¬
note attached a third species to his new genus, namely, M.
Dieteliana ( Melampsorella Dieteliana Sydow). It may be
mentioned in passing that the reason Magnus advanced for
re-distributing the species of Melampsorella was entirely erro¬
neous. Magnus claimed that two sets of rusts were listed
under Melampsorella, one set with stalked uredospores, the
other with catenulate. The fact is that all are stalked. Believ¬
ing that the uredospores of the generic type of Melampsorella
were catenulate, he felt that all those with stalked spores
should be removed and placed in a separate genus by them¬
selves — hence the transfer noted. It was solely on this one
untenable point that his genus Milesina was founded. Fur¬
ther, Magnus’ apology (embodied in the footnote alluded to
above) for discarding Milesia as a name that might be used for
the genus revealed equal lack of knowledge of data essential to
the formulation of a sound pronouncement. He asserted that
as White did not describe the teliospores we do not know what
his rust was — “Ich wahle den Namen Milesina, weil White in
8
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Scot. Naturalist IV p. 162 die Gattung Milesia auf eine Uredo
auf Polypodium vulgare begriindet hat, die vielleicht, wenn
man ihre Teleutosporen kennt, zu Milesina gehort.” . . .
“Meine Gattung Milesina ist aber keineswegs mit der nicht
mehr existenzberechtigten Gattung Milesia White zu identi-
ficieren, da White damit nur den Formcharakter einer unvoll-
kommenen Fruchtform bezeichnet hat, die ebenso gut zur
Uredinopsis, Melampsorella, Melampsoridium etc. gehort und
zu der White oder Berkeley und White einen mir ganz ratsel-
haften Pilz, die Milesia Polygoni B. et White ziehen.” ( Mile¬
sia Polygoni was a misprint for Milesia Polypodii as was ex¬
plained in Grevillia 17:61. 1889). The truth is that White’s
type materials of his species on which the genus Milesia was
founded were carefully preserved and have reposed since 1886
in the Herbarium of the British Museum of Natural History,
along with the correspondence of White, Berkeley and Broome
on them. Moreover, I have found that some of his type ma¬
terials are teliosporic, all of which were available to Magnus.
In fact White’s rust is exactly what he claimed it to be; his
type is extant, and shows that the rust is the same as Sydow’s
Melampsorella Dieteliana. To emphasize still further the
looseness of Magnus’ unauthenticated, apologetic statement,
unhappily to be copied and with even less excuse by Grove
(21), attention needs only to be called to the fact that the rusts
of two of the three genera categorically mentioned by Magnus
are yellow-spored, whereas White took pains to state that the
spores of the rust he described were white. Likewise, the inti¬
mation that Milesia was invalid because founded on the Uredo-
stage (“einer unvollkommenen Fruchtform”) was inaccurate,
because no such authorized rule existed at that time.
Incidentally it may be observed that Magnus primarily
established the genus Milesina on M . Kriegeriana and M.
Feurichii , in the order named. It would follow then that M.
Kriegeriana is the type of the genus Milesina Magnus, with
M. Feurichii as a syntype. Thus it is that Clements and Shear
(l.c.) rightly cite M. Kriegeriana Magnus as the type of Mag¬
nus’ genus Milesina , which they allocate to the position of a
synonym of Milesia White with type M . Polypodii White.
The way is now clear for inquiry into the validity of the
name Milesia in the light of existing, authorized nomenclato-
1932]
FAULL, THE GENUS MILESIA
9
rial rules covering fungi, that is, the rules adopted by the In¬
ternational Botanical Congress of 1910. The rules in
particular that require consideration, are those embodied in
Articles 37 and 49 bis, and Articles 4 and 5.
Article 37 states that “a species or a subdivision of a species,
announced in a work, with a complete specific or varietal name,
but without diagnosis or reference to a former description
under another name, is not valid.” This Article with Article 49
bis therefore determines that any citation of Milesia Poly-
podii White made subsequently to the discovery of the telial
stage by Sydow and his re-naming of the fungus at that time
would not be sufficient to validate the name given by White
unless it were accompanied either by a “diagnosis”, or by a
“reference” to Melampsorella Dieteliana as a synonym. But,
Sydow’s Melampsorella Dieteliana, n. sp., though devoid of
“reference,” escapes the negation imposed by Article 37 be¬
cause it was accompanied by a “diagnosis.”
Article 49 bis defines the conditions that determine the
validity of the names of Fungi. As to the rusts it accords
priority to the first specific name, starting from Persoon’s
Synopsis, applied to the “perfect state,” that is, to the state
which ends in “the teleutospore or its equivalent.” Were no
examples of application appended to Article 49 bis one might
very well interpret the “state” indicated as referring to the
entire diploid state, a state in the rusts that produces both
uredospores and teliospores. These, as is known, are not pro¬
duced by separate mycelial systems, nor in any uniform se¬
quence throughout the group, nor necessarily in separate sori.
Moreover, neither are strictly comparable to the conidia or
“imperfect” spore forms of the Ascomycetes, since the latter
are referable to a haploid “state,” quite a different state from
the one that gives rise to asci. But a reading of the examples
appended seems to indicate that the rule involved was inter¬
preted to mean not the entire diploid state, but that portion of
it embodied in the phrase “teleutospore or its equivalent” — so
commonly and loosely, though conveniently, referred to as the
teleutosporic or teliosporic stage. According to this unquali¬
fied interpretation Milesma Dieteliana (Syd.) Magnus bears
the stamp of validity both as to the name of the genus and the
name of the species.
10
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
There is, however, another aspect of the nomenclatorial
question. The official choice of teliospores as the governing
feature in the priority rule was made, not because teliospores
are the end or “perfect” spore forms, but because of their
supposedly superior taxonomic suitability. It was considered,
and with good reason, that they alone among the various kinds
of rust spores afford in general the most reliable and most
widely applicable means of classification. Now it so happens
that in the course of the extension of our knowledge of rusts, a
genus is revealed in which the teliospores are of extremely in¬
ferior taxonomic utility, while the uredospores are outstand¬
ingly usable. To particularize, I have found few species of
Milesia, out of the twenty-two species and varieties whose telio¬
spores are known, in which the teliosporic characters are of the
slightest taxonomic service in the recognition of species. The
uredospores of Milesia, on the other hand, exhibit unusual, dis¬
tinctive, morphological versatility, so much so that species can
ordinarily be recognized from them alone without reference to
other organs or structures. Practically, in the determination
of species of Milesia, teliospores are usually of minor help,
spermogonia are of secondary importance, aecia are of very
limited service and aeciospores are negligible; on the other
hand uredosporic characters predominantly differentiate the
various species.
That the framers of the accepted Rules anticipated situa¬
tions such as are presented by Milesia, that they recognized
the spirit of the principles underlying the Rules as something
not requiring illogical adherence to defective formulae under
all circumstances, and that they favored or anticipated adapta¬
tions of the Rules if such arose, are more than presumptive
opinions.
Article 4 stresses the importance of “avoidance of all useless
creations of names.” Milesina Magnus constitutes a glaring
example of an unnecessary name, founded on an erroneous
conception and an inexcusable lack of knowledge of attainable
facts. A perfectly good name based on an abundance of
accessible good material was already in existence, so appro¬
priate that the substituted name differs from it only by the
addition of a single meaningless letter. Article 5 recognizes
the weight to be attached to custom. The situation in this
1932]
FAULL, THE GENUS MILESIA
11
connection so far as the particular rusts under consideration
are concerned is illuminating. Of twelve accepted species of
Milesia described in Sydow’s Monographia four only are ac¬
companied by an unembarrassed description of teliospores;
two others mention teliospores but with a question mark; the
remaining six carry the phrase “teleutospores unknown.” The
last set embraces species named by Sydow, Dietel, Jaap and
Wroblewski — named solely from a knowledge of uredinial
characters. Additions of species, some under the name Mile¬
sia, some under Milesina , in which uredospores only were
known, have been made subsequently by Arthur, Dietel, Wine-
land (with approval of Jackson), Cunningham and Faull.
Subjected to unqualified application of Rule 49 bis all of these
names are invalid. On the other hand it has been and con¬
tinues to be the custom to name and describe species of
Milesia on uredinial characters; not a single mycologist — Sy¬
dow, Dietel, Grove, Arthur, etc. — has done other than coun¬
tenance this custom. It could not be otherwise in view of the
facts that the uredosporic characters of this genus are emi¬
nently suited to taxonomic diagnosis, while on the other hand
the teliosporic are in the vast majority of cases entirely value¬
less in separating species and rarely needed in determining the
genus.
Therefore I have accepted Milesia White as the proper name
of the genus and recognize Milesia Polypodii White as its type.
In my judgment sufficient latitude is permitted by the Rules to
sanction these usages, but if not, I anticipate that either the
Rules will be amended so as to extend recognition to uredinial
characters as a criterion of priority, or at the very least that
Milesia White will be listed among the nomina conservanda.
HISTORICAL RESUME
The literature on the rusts of the genus Milesia is compara¬
tively meagre but, scanty as it is, we do find that they have
been studied to some extent from various aspects. Observa¬
tions have been made on their structural features, though these
have been incidental to taxonomic delineations except for more
precise investigations, based on a few American species, of the
uredinia by Moss (38) and the spermogonia by Hunter (27).
Life history studies have been restricted to a single species
12
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
each by Klebahn (32) and Mayor (unpublished), four species
by Kamei (29, 30, 52) and three by Faull (16). Taxonomic
descriptions of twenty-two species in all have been published
under various names; for the main part they have been in¬
complete, mostly of the uredinia only.
The uredinia were the first sori of a Milesia to be observed;
the earliest account of them was given by Fuckel (20) in a de¬
scription of his new species Ascospora Scolopendrii ( Milesia
Scolopendrii) . In this he misinterpreted the uredospores as
“asci” and their granular contents as extremely minute “asco-
spores”; but in general his description of the morphology of
the uredinium was essentially correct. He observed and de¬
scribed the subepidermally located peridium, the short-stalked,
non-catenulate spores (“asci”) and he noted that the latter
were extruded at maturity in white masses or tendrils. His
error concerning the sporal nature of the organs he called
“asci” was soon after corrected by Oudemans (39), though
Oudemans was inclined to regard them as aeeiospores rather
than as uredospores. Fuckel *s studies on “ Ascospora Scolo¬
pendrii” were followed directly thereafter by White’s careful,
independent observations (48) on the uredinia of Milesia
Polypodii White. The uredinia of several other species of
Milesia were subsequently described by various uredinologists;
references to them are embodied in the notes distributed
throughout the taxonomic part of this paper under the sec¬
tions dealing with the relevant species. Except for a brief
digest of the work of Moss, it will suffice here to remark that
entirely divergent views have been held regarding the princi¬
pal uredinial structures of one species or another, for example,
such as to whether the spores are singly attached or catenulate,
whether or not there are paraphyses, whether or not there is a
peridium; and often scant attention was given by earlier ob¬
servers to spore color, that is, as to whether the spores were
white or yellow.
We are indebted most of all to Moss (38) for an accurate
picture of the uredinia of Milesia, as exemplified in M . mar-
ginalis and M. polypodophila, and I have found that his de¬
scription of them for these species applies equally in essential
respects to all other species of Milesia. Moss has demon¬
strated that a peridium is present and has shown that it con-
1932]
FAULL, THE GENUS MILESIA
13
sists of more or less closely adherent terminal, unrelated cells
of primordial columns; he has made clear that these columns
have been misinterpreted as “paraphyses” ; and he has cor¬
rectly interpreted the spores as short-stalked, non-catenulate
derivatives from the basal cells of the primordial columns.
The teliospores, judging from comments on them in the lit¬
erature, have been observed with relative infrequency for most
of the species in which they are known to exist, and for many
of the species not at all. Moreover, they have been imper¬
fectly understood by students of rusts with regard to such
features as the extent of their cellular organization, their mode
of origin, their natural abundance, and their season of occur¬
rence. This is not surprising because, remarkable structures
though they be, they bear little resemblance to spores (Plate
IX, Figs. 34-37), and until one knows when and where to look
for them they are not likely to be seen.
The first teliospores discovered were those of Milesia Krieg-
eriana on fronds of Dryopteris spinulosa. Magnus (35) found
them in the late fall in fronds of the current season. He also
noted basidia growing out from their cells, a single basidium
• from each cell, when the spores germinated. His discovery
was followed by the fortuitous finding of teliospores in a few
other species. But the search for teliospores was so rarely
crowned with success that they were thought to be of rare
occurrence. Sydow’s Monographia expressed the prevailing
belief as to their rarity by the statement that they are formed
“ausserst sparlich.” The explanation for such scant findings is
that search was not made at the proper season. Collectors
expected to find them, just as for M. Kriegeriana, on fronds of
the current season; but I have discovered that M. Kriegeriana
is exceptional with respect to the season at which the telio¬
spores are formed. The teliospores of most species do not
develop in the fall, but instead they develop in great abun¬
dance the following spring on affected overwintered fronds at
about the time Abies is unfolding its leaves. Certainly for the
temperate zone species they are formed in great abundance
and can readily be obtained if sought for at the right time.
The erroneous impression, as expressed by Sydow (40), that
they originate predominantly in the late fall on fronds of the
current season, has continued up to the present to be a deter-
14
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
rent to the study and proper understanding of these rusts, and
in no feature more so than in the determination of their life
histories.
Turning to the aecial stage it is of interest to note that white
peridermia, like those that we now know to be of the Milesian
type, were first described by Peck in America and Kuehn in
Europe. Peck described a form in 1875 from Abies balsamea
under the name Peridermium balsameum, and Kuehn de¬
scribed a similar form in 1884 from Abies alba under the name
Aecidium pseudocolumnare. In neither case were the connec¬
tions with the perfect stages known, nor was it realized that
more than one species with white peridermia might occur on
either Abies balsamea or A. alba. Indeed, the assumption long
prevailed that there was but the one species on Abies alba in
Europe and but one on the various species of Abies in America.
Fraser (Mycologia, 5:233-239. 1913) was the first to demon¬
strate that Peridermium balsameum was passing current as the
name of the aecial stage of several species of Uredinopsis.
Klebahn (32) a few years later showed that Aecidium pseudo¬
columnare included the aecial stage of Milesia Blechni and that
it was probably also the name being given to the aecial stage of
various species of Milesia and Uredinopsis. These conclusions
were confirmed by Kamei in Japan (29, 30 and Annal. Phyto¬
path. Soc. Japan, 2:1-22. 1930), by Faull (16) and by Mayor
(unpublished). Quite naturally the question has now arisen
as to the exact identity of the species from which Peek and
Kuehn drew their descriptions.
To answer this question satisfactorily the type materials
must be available, and criteria for distinguishing in their hap¬
loid phase the species of white-spored rusts on Abies from one
another must be known. So far Klebahn alone has touched the
problem of Aecidium pseudocolumnare and, as we have noted
above, his conclusions are cautious. On the other hand there
have been assumptions, all without verification, as to the iden¬
tity of Peck’s type of Peridermium balsameum. Thus Sydow
(44) refers it to Uredinopsis Strut hiopteridis, etc., and Arthur
(5) to U. americana (U. mirabilis), etc. Concerning the work¬
ing out of a criterional basis for recognizing these rusts on the
Abietinian host Miss Hunter (27) has made substantial prog¬
ress. Her attack has been on the spermogonia, and for the
1932]
FAULL, THE GENUS MILESIA
15
purpose in mind they appear to be favorable organs. In her
comparative study of the spermogonia of the various species of
rusts that attack Abies balsamea she has discovered that Ure-
dinopsis can be distinguished from Milesia by the spermogonia
alone, and that the species of Milesia native to A. balsamea can
likewise be distinguished from one another. Continuing her
investigation she has examined Peck’s collections of Perider-
mium balsOmeum and has found that Peck did not differentiate
among the white-spored peridermia. To him all were P. bal-
sameum. But she has determined that his earliest collections
(unfortunately not dated), one of which was possibly his type
material (though none is so marked), included Milesia inter¬
media. So far then as can be decided from Peck’s herbarial
reliquae Peridermium balsameum may well have been founded
on the O, I stage of M. intermedia — probably so in part at
least.
Life histories have so far been worked out more or less
completely for nine species of Milesia. In all cases species of
Abies have proved to be the alternate hosts. This number
could quite easily be largely increased because teliospores
occur abundantly and they germinate freely. Klebahn (32)
showed that M. Blechni passes to Abies alba and A. cepha-
lonica. Faull (16) has followed through the entire life cycles
of M. intermedia, M. marginalis and M . polypodophila and has
demonstrated that Abies balsamea carries the aecial stage of
all three. Kamei (29, 30, 52) has been successful in similar
studies of four species in Japan, namely, M. exigua, M. Miya-
bei, M. Dryopteridis and M. jezoensis. Finally, Eugene
Mayor informs me that he has derived M. Kriegeriana by sow¬
ing aeciospores of a white-spored Peridermium borne by Abies
alba on one of the common fern hosts of this species.
Relatively, for whatever reason, the discovery of species of
Milesia has proceeded slowly. Magnus (37) in 1909 recog¬
nized three species only; four other rusts known at the time
were transferred subsequently to Milesia, namely, M. Blechni,
M. murariae , M. nervisequa, and M. Scolopendrii. Sydow’s
Monographia (1915) listed twelve valid species, five of which
had been described in the interval. Arthur (2, 4), Kamei
and Hiratsuka (30), Kamei (52), Wineland (28), Faull (16,
18), and Cunningham (9) have since recorded nine more
16
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
(under the name Milesia or Milesina ) as follows: M. consimilis
Arthur, M. australis Arthur, M. jezoensis Kamei and Hirat-
suka, M. Polystichi Wineland, M. Histiopteridis Cunningham,
M. Miyabei Kamei, M. Dryopteridis Kamei, M. polypodophila
(Bell) Faull, and M. exigua Faull. This paper adds fourteen
species and varieties, all new except one transferred from Hya-
lo psora. According to my interpretation, the genus is now
known to comprise thirty-three species and two varieties. In
view of the limited attention that has been given to fern rusts
and the richness of the earth’s fern flora I think it entirely
probable that the genus Milesia includes besides these a large
number of as yet undiscovered species, especially from the
tropics, subtropics and from the mountainous regions of Asia.
HOSTS
It is an exceedingly interesting fact that all successful life
history studies of species of Milesia have shown that their
aecial stage is on the Firs (Abies). The aecial phase of these
rusts on Abies has passed current in Europe, along with that
of various species of Uredinopsis, under the name Aeeidium
(. Peridermium ) pseudocolumnare, and in America under the
name Peridermium balsameum. Of the nine species recorded
below, eight of them are annual on the Firs and restricted to
the needles of the current season; the remaining one, M. poly¬
podophila , is perennial in the stem axes and needles of Abies
balsamea, and its fructifications occur on needles more than
two years of age. Judging from the culture experiments of
Klebahn (32) and Kamei (29) it seems likely that most if not
all of the Firs are susceptible, and that with reference to the
Firs, the species of Milesia are not host restricted. Much ex¬
perimentation, however, bearing on that phenomenon remains
to be done.
The complete list of species of Milesia on the Firs as known
at present follows:
TABLE 1
FIR HOSTS OF SPECIES OF MILESIA
1. Milesia Blechni on Abies cephalonica (Europe).
Milesia Blechni on Abies alba (Europe).
2. Milesia Kriegeriana on Abies alba (Europe).
3. Milesia intermedia on Abies balsamea (North America).
4. Milesia marginalis on Abies balsamea (North America).
1932]
FAULL, THE GENUS MILESIA
17
5. Milesia polypodophila on Abies balsamea (North America).
6. Milesia exigua on Abies firma (Japan).
Milesia exigua on Abies sachalinensis (Japan).
Milesia exigua on Abies Mayriana (Japan).
7. Milesia jezoensis on Abies Mayriana (Japan).
8. M. Miyabei on Abies Mayriana (Japan).
9. M. Dryopteridis on Abies Mayriana (Japan).
Almost equally interesting is the fact that the uredinial and
telial hosts of all species of Milesia are members of the Fili-
cales, and within this Class of the Family Polypodiaceae only.
Lygodium, of the Schizaeaceae, is not the host of any known
species of Milesia, though sometimes cited as such. Distribu¬
tion within the Polypodiaceae is broad, involving representa¬
tives of most of the Subfamilies as shown below in Table 2.
With respect to the question of host restrictions one only (M.
nervisequa, a species of tentative generic allocation) is known
to be intergeneric; moreover, few are known to be interspe¬
cific; the only experimentation in these connections of
which I am aware is recorded in a later section of this mono¬
graph. These experiments and field observations indicate that
the species of Milesia in their relationships to the ferns are
very closely host restricted, a phenomenon the opposite of
that which appears to characterize the aecial generation of
these rusts in relation to the Firs. Especially significant of
the known fern hosts of Milesia is the circumstance that they
“overwinter” their fronds in a more or less green condition.
TABLE 2
DISTRIBUTION OF MILESIA WITHIN THE FAMILY POLYPODIACEAE
•
No. of species and vari-
Subfamily
Genus
eties of Milesia
Woodsieae
Dennstaedtia
1
Aspidieae
Dryopteris
11
Oleandreae
Polystichum
4
Davallieae
Nephrolepis
3
Asplenieae
Blechnum
2
Asplenium
S
Scolopendrium
2
Pterideae
Cryptogramma
1
Histiopteris
1
Pellaea
1
Vittarieae
—
Polypodieae
Acrosticheae
Polypodium
5
The affected fronds so “overwintered” produce a copious crop
of uredospores in the spring before they finally and completely
18
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
die. The result is that species of Milesia appear to be capable
of indefinite perpetuation from year to year on their fern hosts
without the necessity of passing through the aecial phase each
year on the alternate host. The one nearest to exception in
this respect (q.v.) is Milesia intermedia, a species in which
the uredospores are apparently almost eliminated from the
life cycle.
GEOGRAPHICAL DISTRIBUTION
Species of Milesia are to be found in all the major regions of
the world. Nine species have been collected in the United
States and Canada, seven in Central America, West Indies and
South America, eleven in Europe, thirteen in Asia, two in
Africa and one in Australasia. Details have been assembled
for each species and they are incorporated in Table 3.
TABLE 3
GEOGRAPHICAL DISTRIBUTION OF SPECIES OF MILESIA
United States and Canada
A. Eastern United States and Canada
O, I, II, III M. intermedia — Ontario, Quebec, Nova Scotia, Maine, New
Hampshire, Vermont, Massachusetts, New York.
II, III M. fructuosa — Massachusetts, Vermont.
O, I, II, III M. marginalis — Ontario, Quebec, New York, Massachusetts,
New Hampshire.
O, I, II, III M. polypodophila — Ontario, Quebec, Nova Scotia, Maine,
New Hampshire, Vermont, Massachusetts, Connecticut,
New York, Pennsylvania, Tennessee.
B. Western United States and Canada
II, III M. Darken — Oregon, British Columbia.
II, III M. dilatata — Oregon.
II, III M. laeviuscula forma typica — California.
II, III M. laeviuscula forma gylcyrrhiza — Alaska, Washington, Ore¬
gon.
II, III M. Polystichi — Washington, Oregon, California, Idaho, Mon¬
tana.
II, III M. vogesiaca — Oregon.
Central America, West Indies and South America
II M. magellanica — Chili.
II M. australis forma typica — Chili.
II, III M. australis forma irregularis — Ecuador, Colombia, Porto
Rico, Costa Rica.
II M. andina — Ecuador.
II M. columbiensis — Colombia.
II M. Dennstaedtiae — Colombia.
II M. consimilis — Jamaica.
II M. insularis — Porto Rico.
19
1932]
FAULL, THE GENUS MILESIA
Europe
o, i, ii, in
ii, in
ii
ii, hi
i, ii, hi
ii
ii, in
ii, hi
ii, hi
ii, hi
ii, hi
M. Blechni — Great Britain, Germany, France, Switzerland,
Austria,* Czechoslovakia, Italy, Spain, Denmark, Poland.
M. carpatica — Poland, Czechoslovakia.
Mf exigua — Poland.
M. Feurichii — Germany, France, Switzerland, Finland, Czecho¬
slovakia, Hungary.
M. Kriegeriana — Great Britain, Germany, France, Switzerland,
Austria, Denmark, Finland, Poland.
M. Magnusiana — France, Italy, Corsica.
M. murariae — Great Britain, Germany, France, Switzerland,
Austria, Italy, Yugoslavia.
M. Polypodii — Great Britain, Germany, France, Switzerland,
Austria, Spain, Norway, Denmark, Poland, Czechoslovakia.
M. Scolopendrii — Great Britain, Germany, France, Switzer¬
land, Austria, Italy, Poland, Spain.
M. vogesiaca — France, Switzerland, Poland, Czechoslovakia.
M. Whitei — Great Britain, Yugoslavia.
Asia
A. Western Asia
II M. Blechni — Caucasus.
II M. Polypodii — Caucasus.
II M . Scolopendrii — Caucasus.
II M. Whitei — Caucasus.
B. Eastern Asia
II M . Asplenii-incisi — Japan.
II, III M. carpatica var. erythrosora — Japan.
O, I, II, III M. exigua — Japan.
O, (I), II, III M. jezoensis — Japan.
II, III M . Scolopendrii var. sublevis — Japan.
II M. tenuis — Philippine Islands.
II, III M. vogesiaca forma nipponensis — Japan.
O, I, II, III M. Miyabei — Japan.
O, (I), II, III M. Dryopteridis — Japan.
Africa
A. Northern Africa
II M . vogesiaca — Algeria.
B. Southern Africa
II M. nervisequa — Cape of Good Hope, Natal, Transvaal, Mada¬
gascar.
Australasia
II M. Histiopteridis — New Zealand.
* Austria, as it was preceding the war of 1914-18.
Reference to the foregoing table shows that but one species
is recognized in my presentation of geographic distribution as
occurring in both the eastern and the western hemispheres,
namely, Milesia vogesiaca. The rusts in the two hemispheres
diagnosed as this species are morphologically alike; but as
they respectively live on different kinds of hosts they may pos¬
sibly be biological or host-restricted strains, or they may even
be of separate origin. In the latter event they would represent
20
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
a palingenetic species. In any case without fuller knowledge
one would hesitate to venture any speculative deduction from
this one instance of unusual distribution even though ikf . voge-
siaco in America is restricted to the western slope, a region the
flora of which resembles in so many ways the flora of Eurasia.
Still another question, one of much wider scope especially
from the standpoint of the phylogeny of the genus, is suggested
with reference to the tropical and southern hemisphere species,
that is, with reference to Milesian species that occur wholly
beyond the range of Abies. What is the evolutionary history
of these particular species? This question arises, of course,
most appropriately if it be assumed that Abies is the sole host
for the aecial phase of species of Milesia. Unfortunately we
are not yet familiar with the life history of a single one of the
species of Milesia under consideration. Indeed, the only
spores they are known to bear are uredospores except in the
one case of M. australis in Colombia. Because, however, the
teliospores of this species have been found there is reason for
thinking that others of the tropical and subtropical species
produce teliospores also. Considering the complete inconspic¬
uousness of teliospores, the tardiness of their recognition in
the North Temperate Zone, though they abound there, our
slight and wholly casual acquaintance with the rusts referred
to, and our ignorance until recently of the season when telio¬
spores might be found, I am quite inclined to believe that these
rusts, some of them at least, do bear teliospores.
The discovery of teliospores in M. australis was a particu¬
larly happy circumstance. It at once afforded reasonable jus¬
tification for tentatively referring the other similar tropical and
southern hemisphere rusts to the genus Milesia even though
their teliospores have not yet been seen. But more than that,
as bearing on the subject of this discussion, it indicated a perti¬
nent species for a life history investigation. Now, if it should
prove that Abies is the aecial host of M. australis, the assump¬
tion that Abies is the sole host for the aecial phase of species
of Milesia becomes in part an existent fact and more plausible
in its entirety.
Moreover, if Abies is the aecial host of M. australis, and
similarly for the other species found only outside the present
range of Abies, we must accept one of two corollaries, either
1932]
FAULL, THE GENUS MILESIA
21
that the earlier geographical distribution of Abies was very
much greater than at the present time, or that some evolution
of Milesia has taken place on fern hosts without reference to
an aecial stage. Palaeobotany has nothing to offer one way or
the other with respect to the first corollary; palaeobotany in
the tropics and southern hemisphere is a subject still open to
much exploration. As to the second there would be no obstacle
in the matter of perpetuation of species without aeciospores,
for as I point out elsewhere in this paper, all Milesian rusts so
far as have been closely observed propagate from year to year
on their fern hosts from uredospores produced on affected,
“overwintered” fronds. The resulting species in such an evo¬
lutionary sequence would be an imperfect species though not
necessarily lacking the capacity to infect Abies if, as well
might be possible, teliospores are a part of their spore output.
These, in other words, would be species producing uredospores
and teliospores only, or conceivably uredospores only, species
that do not now have and never have had spermogonia and
aecia.
THE TAXONOMY OF MILESIA
My interest in the taxonomy of Milesia began as an outcome
of necessity. In the course of my investigations on the dis¬
eases of Abies it soon became apparent that the utmost confu¬
sion existed with regard to the rusts on Abies. It was discov¬
ered that several rusts on Abies balsamea, different in mor¬
phology and to some extent in behavior, were passing under
the one name Peridermium balsameum. Careful morphologi¬
cal and experimental studies were necessary to untangle them.
One of the first results was the recognition of three hitherto
unknown species of Milesia, and the determination of their
alternate hosts. Fraser had already made valuable contribu¬
tions with regard to Uredinopsis. But in order to secure
authentic materials of the various species for comparison of
their aecia and spermogonia with those of species of Milesia
his work was repeated and then extended so as to unravel the
entire Uredinopsis problem as affecting Abies balsamea. The
same situation was found to exist with respect to western spe¬
cies of Abies, and though it has been impracticable to take up
culture studies in the west, the way has been made clear by
22
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
determining various western species of Milesia on the ferns.
There, as in the east, new species have been discovered, and it
is now made plain that to rest satisfied with guesses as to the
species on Abies is wholly inexcusable. Meanwhile it had be¬
come essential to become acquainted with the species of Mile¬
sia reported from Europe because of suspected misidentifica-
tions on one side or the other, and because such acquaintance
was required in the appraisal of the newly found species.
Similar reasons and curiosity afforded grounds for completing
a survey of the genus.
Very material assistance in the study of the genus resulted
from the discovery of the proper season at which to look for
teliospores. They certainly occur abundantly in the North
Temperate Zone. Referring to Sydow’s Monographia (44) it
is to be noted that in four species only were teliospores un¬
questionably known to exist, and of these in one instance only
was it known at what time they are likely to be found. This
paper records teliospores for twenty-four species and varieties.
MILESIA F. B. White
in Scot. Nat. 4:162 (1877).
Milesina Magnus in Ber. Deutsch. Bot. Ges. 27:325 (1909).
Spermogonia usually hypophyllous, hyaline, deep-seated, glo¬
boid to flask-shaped in vertical section, without ostiolar fila¬
ments, subcuticular or subepidermal.
Aecia hypophyllous, erumpent, peridermial, cylindric, white.
Peridium colorless, rupturing irregularly at apex, inner walls
roughened with warts or short ridges. Aeciospores white,
catenulate, with intercalary cells, globose to ovate or ellipsoid,
verrucose.
Uredinia hypophyllous, rarely petiolicolous, subepidermal,
pustular, or immersed with plane or slightly convex outer sur¬
face, the epidermal covering usually with centrally placed
stomatic pore. Peridium thin, subhemispheric to almost plane,
rupturing by a central, apical pore, sometimes by an irregular
slit; the peridial cells smooth, polygonal, sometimes radially
elongate at sides of the peridium. Uredospores white, borne
singly, short-stalked, obovoid, ellipsoid or subglobose, echinu-
late, verrucose or smooth, pores present but obscure.
1932]
FAULL, THE GENUS MILESIA
23
Teliospores mostly hypophyllous, sometimes amphigenous,
colorless, intraepidermal, rarely one-celled, generally few- to
many-celled, with anticlinal septa, one cell in thickness, one to
a few spores in each containing epidermal cell, rounded, or
more commonly irregular in shape and conforming more or less
to shape of epidermal cell, often completely filling it; walls
smooth, colorless, thin, a single pore in the outer wall of each
cell.
Type species, Milesia Polypodii F. B. White, on Poly podium
vulgare L. Type material carries both uredospores and telio¬
spores; it is deposited in the Herbarium of the British Mu¬
seum of Natural History.
Recent definitions of the genus Milesia are to be found in
Sydow’s Monographia (44) under the name Milesina and in
Arthur’s “Uredinales” in the North American Flora (5) under
the name Milesia. Both are inadequate and in some respects
inaccurate. Thus, as to Sydow’s definition: spermogonia are
no longer unknown, we have knowledge concerning them for
several species; elongated lateral cells are solely transient fea¬
tures in the peridium of the uredinium of most species, and
not seen at all in the peridia of many species at maturity; it is
not true that uredospores are without pores, they can be dem¬
onstrated in many species by boiling the spores in lactic acid;
the uredospores are not always or even characteristically acu¬
leate, in several species they are smooth; teliospores are neither
rare nor sparsely formed. As to Arthur’s definition: the sper¬
mogonia are not always subepidermal, in various species they
are subcuticular; there are no grounds for holding that the
uredinia are dimorphic; dehiscence is not invariably by a cen¬
tral pore, in some instances it occurs by means of an irregular
slit; the same erroneous statement with reference to lateral
cells of the uredinial peridium is made as in Sydow’s defini¬
tion; the teliospores may consist of a large number of cells,
indeed, do consist of a large number in some of the teliospores
of each species, and in no species does the description “2 —
6-celled” adequately apply.
Turning now to descriptions of the species of Milesia it will
be observed that the Key to the Species is based wholly on
uredinial characters. In this Arthur’s incisive opinion (4) as
to “the urediniospores being sufficient to separate the species”
24
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
is largely justified. That they are sufficient as a means of
separating “the genus as well” and that the teliospores are
“wholly negligible taxonomically” are opinions no longer en¬
tirely tenable, as he himself no doubt would now readily grant.
The uredospores are not always generically distinctive; on the
other hand the teliospores are not unimportant in deciding the
generic status of certain species, nor are they entirely devoid
of specific differentiation in every instance in spite of their
amazing uniformity in general.
KEY TO THE SPECIES OF MILESIA
I. Species on Asplenium.
a. Uredinia round to linear, peridial cells loosely coherent in
part.
1. M. Feurichii (on A. septentrionale) . . . . 27
a. Uredinia round, peridial cells closely coherent.
b. Uredospores finely verrucose.
2. M. Asplenii-incisi (on A. incisum ) . 30
b. Uredospores spinulose.
c. Closely and delicately spinulose.
3. M. magellanica (on A. magellcmicum) . . . 31
c. Sparsely and strongly spinulose.
d. Peridial and spore walls thin, 1 [i or less.
4. M. Magnusiana (on A. Adiantum nigrum ) 32
d. Peridial and spore walls much thicker, 1.5-2. 5 p.
5. M. murariae (on A. Rut a muraria ) . 34
II.
III.
Species on Blechnum.
a. Uredospores sparsely and coarsely spinulose.
6. M. Blechni (on B. Spicant ) . 37
a. Uredospores rather closely and delicately spinulose or with
bristles.
b. Spores with delicate spines.
7. M. australis forma typica (on B. auriculatum )
41
b. Spores with spines and bristles.
7. M. australis forma irregularis (on B. occiden¬
tal) ... 41
Species on Cryptogramma.
Uredospores smooth.
8. M. Darken (on C. acrostichoides ) . 46
IV. Species on Dennstaedtia.
Uredospores delicately echinulate.
9. M. Dennstaedtiae (on D. rubiginosa) . . . 48
1932]
FAULL, THE GENUS MILESIA
25
V. Species on Dryopteris. (Also see Addendum, p. 129)
a. Uredinia pustular,
b. Spores smooth.
10. M. dilatata (on D. spinulosa dilatata) . . 49
b. Spores verrucose, subspinulose or roughened.
c. Walls of peridial cells 1.0-1. 3 p thick.
11. M. fructuosa (on D. spinulosa fructuosa , D.
spinulosa americana ) ... 51
c. Walls of peridial cells 2-3 p thick.
12. M. andina (on Dryopteris spec.) . 54
b. Spores spinulose.
c. Spores delicately echinulate, small, 14-28 p long, walls
of spores and peridial cells thin, less than 1 p thick.
13. M. carpatica (on D. Filix mas) . 55
14. M. carpatica var. erythrosora (on D. ery-
throsora) ... 57
c. Spores more strongly echinulate.
d. Spores large, 23-48 p long, walls of spores and
peridial cells thin, about 1 p thick.
15. M. Kriegeriana (on D. spinulosa, D. spinu¬
losa dilatata, D. Filix mas) ... 58
d. Spores smaller, 23-34 p long, strongly echinulate,
walls of spores and peridial cells relatively thick.
16. M. consimilis (on D. patens) . 63
a. Uredinia vesicular (immersed).
b. Spores minutely verrucose, uredinia infrequent and very
inconspicuous.
17. M. intermedia (on D. spinulosa intermedia)
64
b. Spores spinulose, uredinia abundant, evident, though not
conspicuous.
18. M. marginalis (on D. marginalis) . 69
VI. Species on Histiopteris.
Uredospores finely verrucose.
19. M. Histiopteridis (on Histiopteris incisa) 73
VII. Species on Nephrolepis.
a. Uredospores delicately and closely echinulate, narrow, 12-17
P broad.
20. M. tenuis (on N. cordifolia) . 74
a. Uredospores delicately and sparsely echinulate, broader,
14-23 p broad.
b. Intermixture of spiny club-shaped paraphyses.
21 . M. columbiensis (on N. pendula) . 75
b. No paraphyses.
22. M. insular is (on N. rivularis) . 76
26
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
VIII. Species on Pellaea.
Uredospores closely and finely spinulose.
23. M. nervisequa (on P. hastata ) . 77
IX.
X.
Species on Polypodium.
a. Uredospores echinulate.
b. Spores averaging about 16 x 25 p.
23. M. nervisequa (on P. lycopodioides ) ... 77
b. Spores larger, averaging about 19 x 35 p.
24. M. Polypodii (on P. vulgare ) . 81
a. Uredospores finely verrucose, especially at apex.
25. M. jezoensis (on P. vulgare ) . 87
a. Uredospores smooth.
b. Spores attenuate at apex, 38-68 p long, average about
52 p.
26. M. polypodophila (on P. virginianum) . 89
b. Spores not attenuate at apex, 23-48 p long, average
about 33 p.
c. Peridial walls 1-2 p thick.
27. M. laeviuscula forma glycyrrhiza (on P.
glycyrrhiza ) ... 95
c. Peridial walls 2-9 p thick.
27. M. laeviuscula forma typica (on P. calif orni-
cum ) ... 95
Species on Polystichum.
a. Uredospores smooth.
b. Spores small, 18-29 p long, average about 24 p.
28. M. exigua (on P. Braunii, P. japonicum, P.
aculeatum retrorso-paleaceum, P. tripteron )
100
b. Spores larger, 29-43 p long, average about 36 p.
29. M. vogesiaca (on P. lobatum, P. Lonchitis,
P. munitum, P. aculeatum, P. varium) . . 103
a. Uredospores spinulose.
b. Spores strongly echinulate, spores and peridium thick-
walled.
30. M. Polystichi (on P. munitum ) . 108
b. Spores finely echinulate, spores and peridium thin-
walled.
31. M. Whitei (on P. aculeatum ) . Ill
XI.
Species on Scolopendrium.
a. Uredospores coarsely spinulose.
32. M. Scolopendrii (on S', vulgare) . 113
a. Uredospores verruculose or subspinulose.
33. M. Scolopendrii var. sublevis (on S. vulgare)
117
1932]
FAULL, THE GENUS MILESIA
27
I. SPECIES ON ASPLENIUM
I. Milesia Feurichii (Magnus) Faull, n. comb. II and III.
Melampsorella Feurichii Magnus in Ber. Deutsch. Bot. Ges.
20:609 (1902). II and III.
Hyalopsora Feurichii (Magn.) Ed. Fischer in Ured. Schweiz,
p. 475 (1904).
Milesina Feurichii Magnus in Ber. Deutsch. Bot. Ges. 27:325
(1909). II and III.
O and I. Spermogonia and aecia unknown.
II. Uredinia usually petiolicolous, sometimes hypophyl-
lous, and occasionally epiphyllous, subepidermal, scattered on
greenish to brownish areas often involving the blade and the
petiole of the entire frond, round to linear, 0.1 to 0.2 mm.
across and up to 2 mm. long, covered by brownish epidermis
which finally ruptures at a central pore or by a slit; peridium
colorless, typically developed, or in part made up of loosely
coherent cells; peridial cells irregularly polygonal, isodiametric
or somewhat elongate, 7 to 17 p across, with walls 0.5 to 2 p
thick; uredospores colorless, produced singly, with stalks up
to 18 p long, obovoid, ellipsoid or subspherical, 17-26 x 28-
44 p, averaging about 20 x 33 p; spore wall hyaline, 0.5 to
1.5 p thick, rather finely and sparsely echinulate.
III. Telia on overwintered fronds, laminae, rhachides and
stipes, amphigenous, mostly hypophyllous (abaxial), on brown
areas of indefinite extent; teliospores, one to a few each, within
the epidermal cells, frequently in the guard cells, rounded,
usually much elongated or less frequently irregular in outline
and conforming in their narrow, elongated or irregular shapes
to the shape of the containing epidermal cell, sometimes com¬
pletely filling it, one- to fifteen-celled, in the guard cells one-
to four-celled, with anticlinal septa, a single pore in the outer
wall of each cell; the cells of the teliospores with thin, smooth,
colorless walls, 8-19 x 8-27 p; basidia 4-celled, 5-5.5 x 60-80 p;
basidiospores hyaline, globular to broadly obovate, 7-7.5 x
7-9 p.
Hosts and Distribution:
O and I. Unknown.
II. Asplenium septentrionale (L.) Hoffm., in Germany,
Switzerland, Finland, France, Czechoslovakia, Hungary.
28
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
III. Asplenium septentrionale (L.) Hoffm., in Germany,
Czechoslovakia.
Type Locality: Bautzen, Saxony, Germany, on Asplenium sep¬
tentrionale. II and III.
Illustrations: Plate IV, Figs. 13 a-d. — Magnus in Ber.
Deutsch. Bot. Ges. 20: Plate 27, Figs. 1-5 (1902). — Idem 27: Plate
14, Fig. 8 (1909).
Exsiccati: Krieg. Fg. Saxon. 2008, under name Hyalopsora
Feurichii (Magn.) Fischer. — Syd. Ured. 1550, under name Uredo
Scolopendrii (Fuckel) Schroet.
Specimens Examined, all on Asplenium septentrionale. —
Germany: Bautzen, Saxony, July 11, 1900, G. Feurich , Syd. Ured.
1550 (II); type locality. — Dohna, Saxony, Sept. 1902, 1906,
1907, G. Wagner & W. Krieger , Krieg. Fg. Saxon. 2008 (II).
Switzerland: Vallee de Bagues, Valois, Aug. 1906, E. Mayor (II).
France: Plombieres, Vosges, June 1856, Weddell, Herb. L. R. Tul-
asne (II).
Czechoslovakia: Vimperk (Winterberg), July 6, 1932, G. D.
Darker, fronds of current season (II), overwintered fronds
(II, III). Kralovany, July 27, 1932, G . D. Darker (II).
Hungary: Algyogy, April 17, 1916, /. Gydrffy (II).
Milesia Feurichii occurs on the fronds of Asplenium septen¬
trionale in Europe. It has been studied by Magnus (36),
Liro (33), Fischer (19), and Sydow (44), but without agree¬
ment on various points, and especially with regard to the ure-
dinium. Magnus could find no uredinial peridium, and stated
that the peridium was represented by circles of paraphyses.
The spores as he saw them were produced singly and were
short-stalked. He also reported that there were no haustoria.
Liro found a peridium but contended that the spores were
catenulate. Fischer could not locate a peridium, and for this
reason, and without attention to the fact that the spores are
white, transferred the species to Hyalopsora which he wrongly
conceived to be characterized by lack of a peridium. Sydow
saw structures which he interpreted as haustoria; he noted
short cubical cells in the upper part of the uredinium which
he thought represented what might be a more or less eva¬
nescent peridium, and to him the spores were those of a typical
Milesina.
My findings agree with those of Sydow, but with an addi¬
tional comment respecting the peridium. There is a peridium
and no true paraphyses, and the peridium is typical in part at
least. Portions of it, however, may consist of cells that are
1932]
FAULL, THE GENUS MILESIA
29
more or less incoherent; but even they are not fugacious.
With knowledge of the fact that all of the peridial cells of the
uredinium in Milesia are of independent origin, as shown by
the investigations of Moss (38), the unusual, though not
unique, feature of incoherency among some of the cells of the
peridium in M. Feurichii is readily understandable. The periph¬
eral “paraphyses” seen by Magnus were primordial columns
from which spores and peridial cells originate.
A collection of this rust by G. Feurich in Saxony was sent
by him to Magnus under the name Uredinopsis Scolopendrii
(Fckl.) Rostr. Magnus recognized it as an undescribed spe¬
cies and gave to it the name Melampsorella Feurichii; later
(1909) he included it, as a syntype, in his new genus Milesina.
Grove’s claim (23) to the transfer in 1921, under the name
“Milesina Feurichii Grove,” is, of course, an unwarranted
error. Among Feurich’s gatherings was a collection (made in
May 1897 and presumably on overwintered fronds) which,
according to Magnus (36), contained teliospores. There
seems to be no other record of teliosporic material, though such
surely cannot be lacking wherever the rust occurs. A further
examination of the teliospores of M. Feurichii was desirable
because of a statement by Magnus (36) that they occur not
only in the epidermal cells but also “in der subepidermalen
Zellschicht” (inaccurately translated by Sydow (44) as “inter
cellulas subepidermales”). That this might be true seemed
quite possible though I surmised it would be more or less ex¬
ceptional. I have found such a phenomenon in M. marginalis ,
though of relative rarity, and have regarded it as a fortuitous
or a vestigial feature.
Though I have been unable to locate Magnus’ teliosporic
type material, I have finally succeeded in obtaining a collec¬
tion of Asplenium septentrionale bearing the teliospores of M.
Feurichii through the agency of Dr. G. D. Darker. Affected
overwintered fronds collected by Dr. Darker at my request
bear a crop of them as was anticipated, side by side with fronds
of the current season laden with uredinia. It is from this
material that my foregoing description of the teliospores is
drawn. Happily I can now confirm Magnus’ report of telio¬
spores for this species. It should be added that teliospores are
occasionally to be found in the cells of the subepidermal layer,
30
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
but this is of relative infrequency. The typical location of the
teliospores is within the epidermal cells, just as it is in other
species of Milesia.
The teliospores in the epidermal cells of the stipe and rha-
chides are usually, though by no means exclusively, much
elongated, consisting of a single lineal series of cells. This
feature is correlated with the fact that the epidermal layer on
the stipe and rhachides, except in the groove on the abaxial
side, consists of narrow, elongated cells with plane and not
sinuous walls.
An interesting packet of this rust, under the name Hyalop-
sora Feurichii, is preserved in the Cryptogamic Herbarium of
the Paris Museum. I refer to a packet donated by L. R.
Tulasne in 1873. The original label bears the inscription —
“Uredo (Trichobasis) in Asplenio septentrion. (Vosges)
Plombieres. Weddell. Juin, 1856.” It may very well be the
first collection of the genus to be diagnosed as a rust.
2. Milesia Asplenii-incisi Faull, sp. nov. II.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, aggregati vel sparsi, pustulati, rotundati, 0.1-
0.25 mm. diam., peridio ex cellulis hyalinis irregulariter poly-
gonalibus composite cincti; cellulae peridii 8-14 x 8-16 pi,
pariete circa 2 p crasso. Uredosporae obovatae, ellipsoideae
vel rarius subglobosae, remote et subtilissime verruculosae,
hyalinae, 14-17 x 20-26 p, plus minusve 15 x 22 p, episporio
0.7 -1.0 p crasso. Teleutosporae ignotae.
Hab. in foliis Asplenii incisi in Japonia.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, pustular,
round, 0.1 to 0.25 mm. in diameter, covered by cream-colored
epidermis with centrally placed stomatic pore, scattered or
loosely aggregate on pale green or faintly brownish areas of
indefinite extent, often involving entire fronds; peridium hemi¬
spheric, firm; peridial cells hyaline, isodiametrically to irregu¬
larly polygonal, 8-14 x 8-16 p; walls of peridial cells slightly
yellowish, about 2 (j thick; uredospores colorless, short stalked,
obovoid, ellipsoid or occasionally subglobose, 14-17 x 20-26 p,
averaging about 15 x 22 p; walls of spores thin, 0.7 to 1.0 p
thick, with scattered, very fine, dot-like warts.
1932]
FAULL, THE GENUS MILESIA
31
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Asplenium incisum Thunb., in Japan.
III. Unknown.
Type Locality: Inaba-yama near Tottori, Japan, on Asplenium
incisum. II.
Illustrations: Plate VII, Figs. 29 a-d.
Specimens Examined. —
Japan: Inaba-yama near Tottori, March 26, 1930, Naohide Hirat-
suka, on Asplenium incisum (II); part of type.
The uredospores of Milesia Asplenii-incisi, both as to size
and markings, resemble quite closely those of M. intermedia
Faull; but in various other respects the two species are
markedly different and so not readily confused. The uredinia
of the former are abundant, conspicuously pustular and of
medium size; the peridium is firm, and the walls of its cells
comparatively thick. On the other hand the uredinia of M.
intermedia are apparently infrequent, extremely inconspicu¬
ous, and vesicular, practically causing no protuberance of the
overlying epidermis; the peridium is delicate, and the walls
of its cells very thin, being less than 1 m thick. The lesions
caused by these rusts are also dissimilar; those due to M.
intermedia are conspicuous dark-colored spots, while those due
to M. Asplenii-incisi are much diffused and scarcely darker
than the normal unaffected parts. The only material of M.
Asplenii-incisi available to me — a liberal collection received
from Professor Hiratsuka — was gathered in early spring and
showed no teliospores. It is reasonable to anticipate that they
are formed later in the season on the overwintered fronds. It
is also quite likely that Abies will prove to be the alternate
host.
3. Milesia magellanica Faull, sp. nov. II.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregati, pustulati, rotundati, 0.1-
0.2 mm. diam., peridio ex cellulis hyalinis irregulariter poly-
gonalibus composite cincti; cellulae peridii 7-14 m diam.,
pariete 0. 5-1.0 m crasso. Uredosporae obovatae, ellipsoideae
vel subglobosae, brevissime et densiuscule echinulatae, hya-
32
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
linae, 14-20 x 18-30 m, plus minusve 16 x 22 [ j , episporio 0.5-
1.0 n crasso. Teleutosporae ignotae.
Hab. in foliis Asplenii magellanici in Chili.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, pustular, subepidermal,
scattered or loosely aggregate on greenish to brown areas of
indefinite extent, round, 0.1 to 0.2 mm. in diameter, covered by
brown epidermis, which finally ruptures at a centrally placed
stomatic pore, or irregularly; peridium colorless, hemispheric,
delicate; peridial cells irregularly polygonal, isodiametric or
somewhat elongate, 7 to 14 p across, with walls 0.5 to 1.0 p
thick; uredospores white, very short-stalked, obovoid, ellipsoid
or subspherical, 14-20 x 18-30 p, averaging about 16 x 22 p;
spore wall hyaline, 0.5 to 1.0 p thick, with numerous, scat¬
tered, very delicate spines.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Asplenium magellanicum Kaulf., in Chili.
III. Unknown.
Type Locality: Corral, Chili, on Asplenium magellanicum. II.
Illustrations: Plate IV, Figs. 16 a-d.
Specimens Examined. —
Chili: Corral, December 1905, R. Thaxter, on Asplenium magel¬
lanicum (II), type.
The delicate fern Asplenium magellanicum Kaulf. which
carries Milesia magellanica possesses neat, much dissected
fronds, resembling in form and size those of A. Ruta muraria
L. Likewise the general habit of the two rusts which para¬
sitize these ferns, M. magellanica and M . murariae respective¬
ly, is similar, though the two are easily separable as distinct
species.
4. Milesia Magnusiana (Jaap) Faull, n. comb. II.
Milesina Magnus iana Jaap in Fg. Sel. 623, 1913 (nomen) ; in
Verh. Bot. Ver. Brandenb. 57:16 (Jan. 1915). — Sydow,
Monog. Ured. 3:477 (Nov. 1915). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, pustular, subepidermal, scat-
1932]
FAULL, THE GENUS MILESIA
33
tered or loosely aggregate on greenish to brownish areas of in¬
definite extent, round to slightly elongate, 0.1 to 0.4 mm. in
diameter, covered by pale buff-colored epidermis, which finally
ruptures at a centrally placed stomatic pore; peridium color¬
less, hemispheric, delicate; peridial cells elongate and radially
oriented at the sides of the peridium contiguous with the base,
irregularly polygonal, isodiametric to somewhat elongate in
the upper part of the peridium, overlapping, 8-17 \x across,
with walls less than 1 jj thick; uredospores colorless, short-
stalked, obovoid, ellipsoid, rarely club-shaped or subspherical,
17-28 x 28-47 m, averaging about 20 x 35 p; spore wall hyaline,
0.5 to 1.2 m thick, rather strongly and sparsely echinulate, the
spines regularly or irregularly scattered.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Asplenium Adiantum nigrum L., in Corsica, Italy,
France.
III. Unknown.
Type Locality: Ajaccio, Corsica, on Asplenium Adiantum
nigrum. II.
Illustrations: Plate IV, Figs. 15 a-d.
Exsiccati: Jaap Fg. Sel. 623.
Specimens Examined, all on Asplenium Adiantum nigrum. —
Corsica: Ajaccio, Mar. 5, 1913, O. Jaap, Jaap Fg. Sel. 623 (II);
part of type or co-type collection; issued July 1913 under
name uMilesina Magnusiana Jaap n. sp.” without descrip¬
tion.
France: Super-Cannes, Mar. 17, 1932, G. D. Darker (II).
Jaap (Verh. Bot. Ver. Brandenb. 57:16. 1915) informs us
that Milesia Magnusiana occurs in the Ligurian coastal region
of Italy as well as in Corsica. I have not seen specimens from
Italy but have no doubt of the correctness of this statement.
Very recently Dr. G. D. Darker sent me ample material of this
rust collected by himself near Cannes, France, which is close
to the Italian Riviera.
The teliospores have not yet been described. Jaap makes
reference to immature, intercellular ( ! ) teliospores, but with¬
out description. The reference is almost certainly incorrect.
They probably occur abundantly on overwintered fronds about
the middle of spring, and within the epidermal cells.
34
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
5. Milesia murariae (Magnus) Faull, n. comb. II and III.
Uredo Scolopendrii (Fuckel) Schroeter in Cohn, Krypt.-
Flora von Schles. 3(1): 374 (1889), in part. II.
Uredo murariae Magnus in Ber. Deutsch. Bot. Ges. 20:611
(1902). II.
Milesina Murariae (Magnus) Sydow in Monog. Ured. 3:477
(1915). II and III (?). '
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous and petiolicolous, subepidermal,
scattered or loosely aggregate on greenish to brownish areas of
indefinite extent, often involving the entire frond, round, 0.1
to 0.2 mm. in diameter, or elongate (up to 3 mm. long on
petiole or rhachis), covered by light brown epidermis, which
finally ruptures at a centrally placed, stomatic pore or by a
slit; peridium colorless, hemispheric, rather strongly devel¬
oped; peridial cells elongate and radially oriented at the sides
of the peridium contiguous with the base, irregularly poly¬
gonal, isodiametric to somewhat elongate and irregularly dis¬
posed in the upper part of the peridium, 7-15 m across, with
walls 1.5 to 2.0 p thick; uredospores colorless, very short-
stalked, obovoid, ellipsoid or subspherical, 14-23 x 23-37 p,
averaging about 18 by 30 p; spore wall hyaline, 1.5 to 2.5 p
thick, strongly and rather sparsely echinulate.
III. Telia on overwintered fronds, hypophyllous, on in¬
definite, extensive brown areas often involving entire fronds;
teliospores within the epidermal cells, frequently in the guard
cells, hyaline, rounded, or irregular in outline and conforming
to the shape of the containing epidermal cell, often completely
filling it, one- to many-celled (up to fifteen-celled), with anti¬
clinal septa, a single pore in the outer wall of each cell; the
cells of the teliospores with thin, smooth, colorless walls,
irregularly polygonal except along their free margins, 7-16 x
10-25 p.
Teleutosporae intra cellulas epidermidis evolutae, hypo-
phyllae, verticaliter septatae, pluricellulares (1-15 celluli),
leves, hyalinae; cellulae teleutosporarum 7-16 x 10-25 p.
Hab. in foliis Asplenii Rutae murariae in Helvetia.
Hosts and Distribution:
O and I. Unknown.
1932]
FAULL, THE GENUS MILESIA
35
II. Asplenium Rut a muraria L., in Germany, France, Aus¬
tria, Switzerland, Great Britain, Italy, Yugoslavia.
III. Asplenium Ruta muraria L., in Switzerland. Doubt¬
fully reported from France (44).
Type Locality: Not determined. Type locality for teliospores —
Neuchatel, Switzerland, on Asplenium Ruta muraria.
Illustrations: Plate IV, Figs. 14 a-d — W. B. Grove in Jour.
Bot. 59:311 (1921).
Exsiccati: Jaap Fg. Sel. 530. — Rabh. Fg. Eur. 2590, under name
Uredo Scolopendrii (Fuckel) Schroet. — Syd. Ured. 2349 and 2500,
under name Uredo Murariae P. Magn., 2740. — Syd. Myc. Germ.
1163, under name Uredo Murariae P. Magn.
Specimens Examined, all on Asplenium Ruta muraria. —
Germany: Gersfeld, Rhongebirge, July 28, 1906, O. Jaap , Jaap
Fg. Sel. 530 (II). — Gersfeld, Rhongebirge, July 6, 1908, H.
Diedicke, Syd. Ured. 2349 (II). — Freiburg in Baden, 1888,
G. Lagerheim, Herb. N. Y. Bot. Gard., under name Uredo
Scolopendrii Fuckel (II).
France: Forbach, Lorraine, Nov. 10, 1912, A. Ludwig, Syd. Myc.
Germ. 1163 (II).
Switzerland: Montagny, Vaud, Oct. 29, 1911, E. Mayor, Syd.
Ured. 2500 (II). — Montagny, Vaud, July 23, 1919, E. Mayor
(II). — Lausanne, Mar. 30, 1865, L. Favrat, Kew Herb. (II).
— Lausanne, Feb. 23, 1866, L. Favrat, Kew Herb. (II). —
Neuchatel, June 12, 1932, G. D. Darker (II, III); type for
teliospores.
Italy: Bozen, Sept. 4, 1890, P. Magnus, Herb. Bur. PI. Ind., U.
S. Dept, of Agr., Washington, D. C., under name Uredo
Scolopendrii Fuckel (II).
Yugoslavia: Veldes, July 6, 1884, in Mus. Bot. Berol. under name
Gloeosporium Rutae murariae nov. sp.? (inedit), Voss (II).
Uredinologists have long been familiar with Milesia mura¬
riae, but from its uredinia only. The uredinia are conspicu¬
ous and the uredospores are produced abundantly. Magnus
(36) was right in recognizing it as a distinct species though his
basis of separation was wrong. He at first thought it to be the
same as Melampsorella Feurichii Magnus [Milesia Feurichii
(Magnus) Faull] but after finding a peridium in M. murariae
and believing that there was none in M. Feurichii he decided
to describe the former as a new species, Uredo murariae.
Magnus described the uredinia in quite accurate detail, espe¬
cially with respect to the peridia. He observed that the
peridium is small-celled, the cells irregularly disposed in a
single layer, and not in longitudinal rows, and that it opens by
a centrally placed, apical pore located immediately under an
36
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
epidermal stoma which is enlarged as a result of the growth
of the uredinium. He further noted that the peridial cells
appear to be apical terminal cells cut off from palisade-like
cells, several sets or layers of which surround the uredinium.
Though it escaped him that these palisade cells also give rise
to the spores, his observations as to the origin of the peridial
cells have been confirmed by Moss (38) and others.
Grove (23) redescribed the uredinial stage of this rust in
1921 under the title “Milesina murariae, comb, nov.”, appar¬
ently unaware that that combination had been made several
years previously by Sydow (44). He remarked that “this
very rare parasite has been found in Scotland before (Crypt.
Soc. Report, 1907), but was at that time included under the
old Milesia Polypodii B. White.” He observed that the spores
were colorless. His illustrations are poor, and his descriptions
of spore markings and of the uredinial envelope are inaccurate.
The uredinia occur profusely on affected fronds, on stipes
and rhachides as well as on the blades. They resume their
activity in early spring on overwintered fronds, and discharge
large quantities of viable spores at about the time the new
fronds are unfolding. The latter are promptly infected and
soon thereafter they, too, bear numerous uredinia from which
emerge tendrils and masses of pure white uredospores. Thus
for a few weeks in the early part of the growing season, before
the old fronds have completely died, one may find active ure¬
dinia on both the old and the new fronds of the same plant.
The uredospores are strongly echinulate, but the distribu¬
tion of the spines is not always uniform. Occasionally one finds
spores on which there are smooth areas of variable and some¬
times considerable extent. The spines, too, are variable in
height, measuring up to 2.2 jj.
Long as M. murariae has been known there has been com¬
plete uncertainty up to the present regarding its teliospores.
Magnus (36) informs us that in one specimen only, collected
in November, did he find what might be teliospores — “in ein-
zelnen Epidermiszellen hyaline, Melampsorella-artige Zellbil-
dungen.” The Sydows (44) recount a similar experience.
They diligently searched for teliospores in excellent material
sent to them at various times throughout the year, but in one
frond only, collected towards the end of November, did they
1932]
FAULL, THE GENUS MILESIA
37
find anything more than a few intracellular spores, never more
than 2 -celled, which they regarded as immature teliospores.
The interpretation in each instance is improbable. The telio¬
spores of M. murariae, as a matter of fact, are neither excep¬
tional nor rare; they are produced, just as in other species of
Milesia in which they are known, in abundance. But to find
them the search must be made, not on fronds of the current
season, but on affected overwintered fronds in the spring.
Such fronds, recently collected by Dr. G. D. Darker at my re¬
quest, carry, as was anticipated, a copious crop of them. It is
from this material that my foregoing description of the telio¬
spores has been drawn.
The next step in the study of M. murariae is to work out its
life history. Presumably Abies is the alternate host.
II. SPECIES ON BLECHNUM
6. Milesia Blechni (Syd.) Arthur in Bot. Gaz. 73:61 (1922),
in part. (II). O, I, II and III.
Uredo Scolopendrii (Fuckel) Schroeter in Cohn, Krypt.-
Flora von Schles. Band 3(1):374 (1889), in part. II.
Uredinopsis Scolopendrii (Fuckel) Rostrup in Bot. Tidsskr.
21:42 (1897), in part. II.
Melampsorella Blechni Sydow in Annal. Mycol. 1:537 (1903).
II and III.
Milesina Blechni Sydow in Annal. Mycol. 8:491 (1910). II
and III.
Aecidium pseud ocolumnare J. Kuehn in Hedwigia 23:168
(1884), (acc. to various authors), in part. — See Klebahn
(32).
O. Spermogonia on needles of current season, amphi-
genous, mostly hypophyllous, immersed, more or less flask-
shaped in sectional view, apparently subepidermal in origin,
covered by epidermal cells the inner walls of which become
more or less disorganized, 110 to 175 p broad by 105 to 150 p
deep; spermatia hyaline, narrowly cylindrical, 4-6 p long.
I. Aecia hypophyllous on needles of current season, in two
rows, one on each side of midrib, white, cylindrical, 0.3 to 0.4
mm. in diameter; peridium colorless, delicate, rupturing at the
apex; peridial cells polygonal, elongated vertically, overlap¬
ping, in a single layer, 12-24 x 24-40 p, with outer walls
smooth, about 1 p thick, with inner walls finely and densely
38
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
waited, the warts often in irregular lines, 2. 5 -3.0 p thick;
aeciospores ellipsoid, ovoid or globose, mostly elongated,
white, 21-27 x 27-36 p, densely and rather coarsely warted ex¬
cept on one side, on which the warts are minute; wall color¬
less, thin.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite ex¬
tent, pustular, 0.17 to 0.4 mm. across, rupturing at a centrally
placed stoma of the overlying epidermis; peridium hemi¬
spheric, hyaline; upper peridial cells isodiametrically to irreg¬
ularly polygonal, 7-12 p across, overlapped; lateral peridial
cells radially elongated; walls of peridial cells hyaline, thin;
uredospores colorless, numerous, short-stalked, obovoid to
ellipsoid, 15-23 x 26-45 p, averaging about 19 x 33 p; stalks
of uredospores up to 12 p long by 6 p wide; walls of spores
thin, 0.7 to 1.0 p thick, with scattered, rather coarse echinu-
lations.
III. Telia on overwintered fronds, on indefinite, extensive,
brown areas, at times involving entire pinnae; teliospores with¬
in the epidermal cells, frequently within even the guard cells,
abundant in the lower epidermis, occasional in the upper epi¬
dermis, hyaline, rounded, or irregular in outline and conform¬
ing to the shape of the containing epidermal cell, at times com¬
pletely filling it, one- to seventy-celled, with anticlinal septa;
spores in the guard cells up to twelve-celled; the cells of the
teliospores with thin, smooth, colorless walls, irregularly polyg¬
onal except along their free margins, 6-11 x 8-16 p.
Hosts and Distribution:
O and I. Abies alba Mill, and A. cephalonica Loud., in
Germany, according to Klebahn (32). This stage is Aecidium
pseudocolumnare J. Kuehn, in part ; the peridermia of all white-
spored rusts on Abies in Europe have probably been comprised
under this name, just as in America they have passed under
the name of Peridermium balsameum Pk.
II. Blechnum Spicant (L.) With., in Germany, Austria,
Switzerland, France, Great Britain, Denmark, Poland, Spain,
Czechoslovakia, Caucasus.
III. Blechnum Spicant (L.) With., in Germany, France.
Type Locality: Grosser Winterberg, Schmilka, Saxony, Ger¬
many, on Blechnum Spicant. II and III.
1932]
FAULL, THE GENUS MILESIA
39
Illustrations: Plate II, Figs. 5 a-d. — Grove in British Rust
Fungi, Fig. 282, p. 377 (1913). — R. Gonzalez Fragoso in Flora
Iberica, Uredales, II, Fig. 137, p. 281 (1925).
Exsiccati: Syd. Ured. 1841 and 2092, under name Melampsorella
Bleckni Syd., 2739. — Syd. Myc. Germ. 61 and 311, under name
Melampsorella Blechni Syd., 877. — Jaap Fg. Sel. 527 and 774 a,
b. — Schroet. Pilze Schles. 128, under name Uredo Scolopendrii
(Fuckel) Schroet. — Krieg. Fg. Saxon. 816, under name Uredo Sco¬
lopendrii (Fuckel) Schroet. — Petrak Flora Bohemiae et Moraviae
Serie II Abt. 1 Pilze 1732. — Vestergren Microm. Rar. Sel. 692, under
name Uredo Scolopendrii (Fuckel) Schroet.
Specimens Examined. O, I on Abies alba and A. cephalonica;
II, III on Blecknum Spicant. —
Germany: Schmilka, Saxony, Aug. 28, 1903, H. and P. Sydow, Syd.
Ured. 1841 (II); type locality. — Winterberge, July and Au¬
gust 1890, 1891 and 1892, G. Wagner, Krieg. Fg. Saxon. 816
(II). — Klebahn’s cultures, July 25, 1915, H. Klebakn, Jaap
Fg. Sel. 774 a (O, I). — Rehberger Graben, Harz, Aug. 24,
1904, P. Sydow, Syd. Myc. Germ. 311 (II). — Feldberg,
Baden, July 25, 1903, G. Lagerkeim, Vestergren Microm.
Rar. Sel. 692 (II). — Stiitzerbach, Thuringia, July 20, 1911,
O. Jaap, Jaap Fg. Sel. 527 (II). — Schwarzwald, near St.
Georgen, Sept. 1913, P. Sydow, Syd. Ured. 2739 (II). —
Klecker Wald, Hanover, May 12, 1915, on overwintered
fronds, O. Jaap, Jaap Fg. Sel. 774 b (II, III). — Einsiedelei,
Westfalen, Aug. 31, 1919, A. Ludwig, Herb. Bur. PI. Ind.,
U. S. Dept, of Agr., Washington, D. C. (II).
Switzerland: Hasleberg, Bern, Sept. 3, 1906, Ed. Fiscker, Syd.
Ured. 2092 (II). — Saignolis, Neuchatel, July 11, 1929, E.
Mayor (II).
France: Hohneck, Central Vosges, July 16, 1910, H. Sydow, Syd.
Myc. Germ. 877 (II). — Col de Jau, Aude, Apr. 29, 1932.
G. D. Darker and G. Fenwick-Owen (II, III). — Ax-les-
Thermes, Ariege, May 6, 1932, G. D. Darker (II).
England: Lytchett Minster, Dorset Co., Nov. 26, 1931, G. D.
Darker (II). — Brockenhurst, Hants Co., Nov. 25, 1931,
G. D. Darker (II).
Poland: Sokolskie, eastern Carpathians, June 1914, A. Wroblewski
(II). — Mount Tatra, July 10, 1916, A. Wroblewski (II). —
Leduscul, eastern Carpathians, March 1916, A. Wroblewski
(II).
Czechoslovakia: Hochgesenke, Gr. Kessel, Mar. 19, 1923, F.
Petrak, Petrak Flora Bohemiae et Moraviae II Serie Abt. 1
Pilze 1732 (II).
Spain: Llanes (Oviedo), Sept. 1920, P. Unamuno, Herb. Bur. PI.
Ind., U. S. Dept, of Agr., Washington, D. C. (II).
Caucasus: Batum, Sept. 1917, W. Siemaszko, Herb. Bur. PI. Ind.,
U. S. Dept, of Agr., Washington, D. C. (II).
40
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Two descriptions of the uredo stage of Milesia Blechni ante¬
date Sydow’s (42); Sydow makes reference to neither. The
first was by Schroeter (41) in his account of Uredo Scolo-
pendrii (Fuckel) Schroet., n. comb., as represented by Silesian
collections on Asplenium Ruta muraria and Blechnum Spicant
— rusts subsequently segregated as Milesia murariae and M.
Blechni respectively. He correctly appraised the spores he
observed as uredospores and made special note of the fact that
they are white as seen in tendril-like extruded masses. The
second description was by Rostrup (40) and it was note¬
worthy as indicative of accurate observation. Referring to a
rust found in Jutland on Blechnum Spicant he observes (trans¬
lating freely) that “the parasitized fronds were decolorized in
spots and on the lower surface of these spots were brown pus¬
tules from which chalk- white spores emerged in strings or ten¬
drils.” “As to this fungus on B. Spicant — Fuckel described a
white-spored fungus on Scolopendrium vulgare under the name
Ascospora Scolopendrii, taking it to be a Pyrenomycete. He
mistook the spores for asci, thinking that they were filled with
extremely minute spores. Oudemans placed the same fungus
in the Uredineae, but regarded its fructification as an aeci-
dium. Schroeter with better reason called it Uredo Scolo¬
pendrii; in Silesia it was found on Asplenium Ruta muraria
and on Blechnum Spicant. After considering all features of
the fungus there is no doubt it should be called Uredinopsis
Scolopendrii (Fuckel) Rostr., n. comb.” — a combination to be
adopted later by Fischer (19).
Milesia Blechni is known from Europe and the Caucasus
only, but since its host occurs also widely spread in Asia and
along the Pacific slope of North America from Alaska to Cali¬
fornia, quite possibly the range of this rust is much greater
than has been so far reported. It would be of special interest
to know whether or not it is represented on B. Spicant in
America. Two other species of Blechnum in tropical parts of
America are known to carry a Milesia, but this is a different
rust, M. australis, and not M. Blechni, as has frequently been
wrongly assumed. I would point out that they differ in ure-
dinial characters, and also that the cells of the teliospores of
M. Blechni are considerably smaller than are those of M. aus¬
tralis forma irregularis.
1932]
FAULL, THE GENUS MILESIA
41
Quite recently Arthur (4) on the basis of uredo characters
has again raised the question as to whether or not M. Blechni
is the same as M. Scolopendrii and he proposes to include both
under the latter name. It is true that the uredospores and the
uredinia of the two species are quite similar ; but I have found
that the uredospores of M. Scolopendrii are consistently larger
than those of M. Blechni , and in general their walls are thick¬
er; likewise the cells of the teliospores are smaller than are
those of M. Scolopendrii. These rusts, too, are probably host
restricted and it is not unlikely that their spermogonia will be
found to be specifically distinguishable.
Up until 1931 M. Blechni was the only European species of
Milesia in which the life history had been followed throughout
(Klebahn, 32). Recently, however, Dr. Eugene Mayor in¬
formed me that he has now succeeded in establishing the con¬
nection of M . Kriegeriana on Dryopteris spinulosa with a white
peridermial rust on Abies alba. It is of particular interest to
discover in these two rusts, as in the species of the same genus
studied by Hunter (27), that their spermogonia are of specific
diagnostic significance. Here again is evidence that within the
genus Milesia the uredinia and the spermogonia have in gen¬
eral responded more strikingly than the other organs to the
play of evolutionary forces.
7. Milesia australis Arthur in Bull. Torrey Bot. Club, 51:53
(1924). (II). II and III.
Uredo Blechni Dietel & Neger in Bot. Jahrb. 22:358 (1897).
Milesina Blechni Syd., in part, acc. to P. Dietel in E. Mayor’s
“Contribution a l’etude des Uredinees de Colombie.” Mem.
Soc. Sci. Nat. Neuchatel, 5(2):557 (1914). II and III.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely aggregate on greenish to brownish areas of indefinite
extent, pustular, round, 0.1 to 0.3 mm. in diameter, rupturing
at a centrally placed stoma in the overlying epidermis; peridi-
um hemispheric, hyaline; upper peridial cells isodiametrically
to irregularly polygonal, 8 to 16 p across; lateral peridial cells
radially elongate; walls of peridial cells hyaline, thin, about
1 p thick; uredospores colorless, very short-stalked, variable
and often irregular in shape, obovoid, ellipsoid or subglobose,
42
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
14-23 x 22-40 p, averaging about 18 x 30 [i; walls of spores
0.8- 1.5 p thick, with rather numerous, small, delicate echinu-
lations, or in some collections with irregularly scattered bristles
or spines up to 2 p long, short spines and minute warts.
III. Telia hypophyllous, on indefinite, extensive, brown
areas; teliospores within the epidermal cells, common in the
guard cells, hyaline, rounded, or irregular in outline and often
conforming to the shape of the containing epidermal cell, at
times completely filling it, one- to many-celled, up to thirty or
more, with anticlinal septa; the cells of the teliospores with
thin, smooth, colorless walls, irregularly polygonal except along
their free margins, 8-19 x 11-27 p.
Hosts and Distribution:
O and I. Unknown.
II. Blechnum auriculatum Cav. ( B . hastatum Kaulf.), in
Chili. ( M . australis forma typica).
Blechnum occidentale L., in Colombia, Ecuador, Porto
Rico, Costa Rica. ( M . australis forma irregularis) .
III. Blechnum occidentale L., in Colombia.
Type Locality: Forma typica. Concepcion, Chili, on Blechnum
auriculatum. II. (Not Concepcion, Peru, as stated by Arthur, 1925,
and by Kern and Whetzel, 1926).
Forma irregularis. Central Andes near Medellin, Antioquia, Co¬
lombia, on Blechnum occidentale. II and III.
Illustrations: Milesia australis typica. Plate II, Figs. 6 a-d. —
Milesia australis irregularis. Plate II, Figs. 7 a-d. — Dietel in Mem.
Soc. Sci. Nat. Neuchatel 5(2) : 5 5 7 Figs. 65 a, b (1914).
Exsiccati: Syd. Ured. 44 a, under name Uredo Blechni Diet. &
Neg., n. sp.
Specimens Examined. —
A. Milesia australis forma typica on Blechnum auricu¬
latum.
Chili: Concepcion, May 1896, F. W. Neger, Syd. Ured. 44 a (II);
presumably part of type. — Corral, Dec. 1905, R. Thaxter
(H).
B. Milesia australis forma irregularis on Blechnum occi¬
dentale.
Colombia: Central Andes, alt. 1700 m., near Medellin, Antioquia,
Aug. 16, 1910, E. Mayor, Farlow Herb. (II, III); type. —
Central Andes near Medellin, Antioquia, Aug. 14, 1910, E.
Mayor (182 c) (II).
Ecuador: 1910, G. Lagerheim, Riksmuseets Botaniska Avdelning,
Stockholm (II).
Porto Rico: Rio Prieto, June 22, 1924, H. H. Whetzel, F. D. Kern
1932]
FAULL, THE GENUS MILES I A
43
and R. A. Toro (2296), Herb. Explorations of Porto Rico,
Herb. N. Y. State Coll, of Agr. (II).
Costa Rica: San Jose, Dec. 12 and Dec. 19, 1924, H. Sydow, Far-
low Herb., under name Milesina Blechni (II).
The type of Milesia australis was described by Dietel and
Neger (13) from uredo material collected in Chili, on Blech-
num auriculatum ( B . hastatum) . They stated that it differed
from uUredinopsis Blechni” (that is, Milesia Blechni ) by its
smaller spores and more delicate echinulations.
As indicated above I have examined two collections of Mile¬
sia australis from Chili, both recorded as occurring on Blech-
num auriculatum, and five collections from three widely sep¬
arated areas in the Caribbean and northwestern South Ameri¬
can regions, all recorded as occurring on Blechnum occiden¬
tal. In general the rusts in these various collections are very
similar in their morphological features. The greatest varia¬
tion is shown in Mayor’s collections from Colombia and Lager -
heim’s from Ecuador, but only with respect to the spinulations
on the uredospores; these are irregularly scattered spines up
to 2 n long (Plate II, Figs. 7 a-d) more or less aggregated at
the poles, short spines and minute warts, and frequently there
are smooth areas of varying extent. If these features were
constant there would be justification in recognizing the Colom¬
bian and Ecuadorian rust as a distinct variety at least. But
an examination of the Porto Rican and the Costa Rican collec¬
tions on what is claimed to be the same specific host reveals
spores that are more or less free from bristles, and some of
them are as evenly spinulose as the Chilean plants. Under the
circumstances it seems wisest to tentatively recognize all as
one and the same species. It is convenient, however, to desig¬
nate the rust on B. auriculatum as forma typica Faull and the
one on B. occidentale as forma irregularis Faull, f. nov.
The following table records the measurements of 40 uredo¬
spores taken at random from each of five collections:
TABLE 4
Range A verage
1. M. australis typica (Concepcion, Chili)
2. M. (kistralis typica (Corral, Chili)
3. M. australis irregularis (Colombia)
4. M. australis irregularis (Costa Rica)
5. M. australis irregularis (Porto Rico)
14-23 x 23-40 18 x 32
14-20 x 23-32 17 x 27
14-23 x 23-37 18 x 29
14-23 x 22-39 18 x 30
14-23 x 25-40 \i 18 x 31 n
44
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Referring to M. australis forma typica, under the name
Uredo Blechni Diet. & Neger, Sydow (Monogr. Ured. 4:552.
1924) expressed the view that it is a Milesina, but, re-echoing
Dietel and Neger (13), because of the smaller size and more
delicate spinulation of its spores it could not be M. Blechni.
With these views I am fully in accord. Arthur (4, 5) likewise
agrees, thus reversing an earlier opinion (3), but he errs in
also including under the name M. australis a rust on Lygo-
dium polymorphum from Salvador; the uredospores of the lat¬
ter are like those of a collection from Brazil on L. polymor¬
phum which Arthur himself (4) has demonstrated as quite
removed from a Milesia.
It is needless to add that M. australis forma irregularis is
equally different from M. Blechni; but particular interest at¬
taches to one of Mayor’s collections of this rust from Colom¬
bia because of the fact that it carries teliospores, the only
known instance of the telial stage in species of Milesia from
tropical or subtropical regions. Dietel (12) states that he
found an abundance of teliospores in that collection, and his
careful, detailed descriptions of them are supplemented by
good drawings. I have been equally fortunate in finding abun¬
dant teliospores in a portion of Mayor’s collection found in the
Farlow Herbarium, and it is on this material that my study of
the telial stage is based. I have been able to corroborate and
extend Dietel’s observations. It is to be noted that the cells
of the teliospores are in general larger than those of M. Blechni
(q.v.). The finding of teliospores in M. australis leads one to
anticipate that the telial stages of other tropical and subtrop¬
ical species will be found in collections made at the proper sea¬
son. They should be sought in old, rusted fronds, both at the
close of their growing season and early in the subsequent grow¬
ing season. It has long been supposed that the teliospores of
the temperate zone species are rare, but in my experience that
is not the case. Actually they occur abundantly. The same
thing may be true of several, if not all, of the tropical or sub¬
tropical species.
Inquiry as to the aecial stage of the tropical and subtropical
species of Milesia naturally follows. Enough experimentation
has been carried out with temperate zone species to lead one to
expect Abies to be the carrier of this stage. As Abies is re-
1932]
FAULL, THE GENUS MILESIA
45
stricted to the North Temperate Zone, however, it is certain
that the species of Milesia in such regions as New Zealand, the
Philippine Islands, and South America persist without refer¬
ence to Abies. Whatever may be said regarding the hetero-
ecism of the tropical and subtropical species of Milesia is pure
speculation. Yet with some show of reason one may surmise
that an aecial host is not necessary to their perpetuation. We
know that temperate zone species may continue from year to
year on the fern hosts without an alternate phase on Abies be¬
cause of the fact that uredospores develop abundantly on
rusted, overwintered fronds; a similar phenomenon may char¬
acterize the other species, and be their only means of perpetua¬
tion. More speculative is the supposition that the tropical and
subtropical species of Milesia have never had an aecial stage
in their existing specific forms; they may be evolutions from
the generation on the fern host solely. They may be imperfect
rusts of a distinctive and thus far unrecognized type. If
they have been evolved from long cycle species with Abies as
one of their hosts, it is of course by no means certain that
basidiospores if formed from their teliospores have retained
the capacity to infect Abies. It would now be of great interest
to test this matter in such a species as M. australis. If suc¬
cessful the next query would be as to whether or not there had
been a world-wide distribution of Abies at some eras in the
past, or if the evolution of Milesia beyond the range of Abies
has been as suggested above. If not successful there would
remain the possibility of some other alternate host; its discov¬
ery should not present an especially difficult task.
Kern and Whetzel, in an account of their collections of M.
australis on Blechnum occidentale in Porto Rico (Frank D.
Kern and H. H. Whetzel, Some New and Interesting Porto
Rican Rusts. Mycologia, 18:39-47. 1926) took occasion to
cite a rust on a Porto Rican collection of Polytaenium lanceo-
latum (L.) Benedict ( Antrophyum lanceolatum Kaulf.) from
a Sintenis specimen, no. 1800, Herbarium N. Y. Botanical
Garden, as also M. australis. But the rust on P. lanceolatum
is something else entirely; apparently it is an undescribed spe¬
cies. Its uredinia are provided with a peridium, they are
amphigenous, mostly epiphyllous, and relatively large; the
uredospores are also relatively voluminous, averaging 28 x 38
46
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
M, varying in width from 23 to 31 \a. It is impossible to rec¬
ognize the color of the spores from the dried specimens; they
may very well be yellow in color when freshly extruded.
III. SPECIES ON CRYPTOGRAMMA
8. Milesia Darken Faull, sp. nov. II et III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli
et petiolicoli, epidermide tecti, sparsi vel aggregate pustulati,
rotundati, 0.2-0. 3 mm. diam., peridio ex cellulis hyalinis irregu-
lariter polygonalibus composito cincti; cellulae peridii 8-14 p
diam., pariete 1. 5-2.0 m crasso. Uredosporae obovatae vel
ellipsoideae, leves, hyalinae, 15-20 x 36-64 \a, plus minusve
17 x 50 n, episporio circa 1 m crasso. Teleutosporae in hypo-
phyllo et rarius in epiphyllo intra ceJlulas epidermidis evolu-
tae, verticaliter septatae, pluricellulares, leves, hyalinae; cel-
luli teleutosporarum 8-21 x 11-25 m*
Hab. in foliis Cryptogrammes acrostichoidis in Oregon et
British Colombia, America bor.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, also on the petiole, subepider-
mal, scattered or loosely grouped on greenish to brown areas
of indefinite extent, sometimes involving entire fronds, pustu¬
lar, round, 0. 2-0.3 mm. in diameter, covered by a brownish dis¬
colored epidermis which finally opens by a central pore; peridi-
um hemispheric, firm; peridial cells hyaline, isodiametrically
to irregularly polygonal, 8-14 jj across; walls of peridial cells
hyaline, 1.5 -2.0 m thick; uredospores colorless, white in mass,
numerous, produced singly, short-stalked, obovoid to ellipsoid,
gracefully asymmetric as a rule, 15-20 x 36-64 p, averaging
about 17 x 50 m; walls of spores thin, about 1 m thick, smooth.
III. Telia on overwintered fronds, hypophyllous and occa¬
sionally epiphyllous, on indefinite, extensive, brown areas, at
times involving entire fronds; teliospores within the epidermal
cells, sometimes even within the guard cells, hyaline, rounded,
or irregular in outline and conforming to the shape of the con¬
taining epidermal cell, often completely filling it, one- to many-
celled, up to twenty-five, with anticlinal septa, a single pore in
the outer wall of each cell; the cells of the teliospores with
1932]
FAULL, THE GENUS MILESIA
47
thin, smooth, colorless walls, irregularly polygonal except along
their free margins, 8-21 x 11-25 p.
Hosts and Distribution:
O and I. Unknown.
II and III. Cryptogramma acrostichoides R. Br., in Ore¬
gon, British Colombia.
Type Locality: Mouth of Mud Creek, Clackamas Co., Oregon,
on Cryptogramma acrostichoides. II and III.
Illustrations: Plate VI, Figs. 24 a-d.
Specimens Examined, all on Cryptogramma acrostichoides. —
Oregon: Junction Mud Creek and Salmon R., July 12, 1929, G. D.
Darker (2759), Arnold Arb. Path. Herb. 254 (II, III), telia
on overwintered fronds; type. — Junction Mud Creek and
Salmon R., July 19, 1929, G. D. Darker (2830), Arnold Arb.
Path. Herb. 325 (II, III), telia on overwintered fronds. —
Yokum Falls, Government Camp, July 15, 1929, G. D.
Darker (2784), Arnold Arb. Path. Herb. 279 (II, III), telia
on overwintered fronds. — Yokum Falls, Government Camp,
Sept. 7, 1931, /. H. Fault (II).
British Colombia: Daisy Lake, April 28, 1932, T. W. Childs (II,
III).
This distinctive Milesia is the first one recorded as occur¬
ring on a Cryptogramma. So far we know the diploid stage
only, but on the fern host the uredinia and the telia were
found in abundance.
The uredinia are typical for the genus. Each uredinium de¬
velops immediately below a stoma, and this serves as the point
at which the “central pore” is developed. When spore dis¬
charge begins the peridial cells just under the stoma are
crushed and broken outwards by the emerging spores, the inner
lateral walls of the guard cells are disorganized and their over-
lying cuticle ruptured. The peridium readily lends itself to
study because it can be quite easily freed in part from the
overlying epidermis. The suggestion of a circle of paraphyses,
mistakenly interpreted as such in various other fern rusts, is
evident in young uredinia. The elements in question are
plainly primordial columns from which spore mother cells and
peridial cells proper have not yet been fully organized. In
more mature uredinia these wholly or largely disappear and
the peridium is seen to consist of polygonal cells without a
marginal supplement of elongated structures.
The telia are abundant on overwintered fronds; hence little
48
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
trouble should be experienced in obtaining materials for cul¬
ture experiments. One awaits with interest now a determina¬
tion of the complete life cycle. While M. Darken is well
equipped to propagate itself indefinitely on the fern host, be¬
cause of prolific uredospore reproduction on overwintered
fronds, there can be little question that the peridermial stage
will be found to occur on Abies.
IV. SPECIES ON DENNSTAEDTIA
9. Milesia Dennstaedtiae (Dietel) Faull, n. comb. II.
Milesina Dennstaedtiae Dietel in E. Mayor in Mem. Soc. Sci.
Nat. Neuchatel, 5(2):558 (1913). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite ex¬
tent, round or slightly elongate, about 0.1 mm. in diameter,
covered by dark brown epidermis which finally ruptures at a
centrally placed stomatic pore; peridium colorless, delicate;
upper peridial cells irregularly polygonal, 8 to 16 m across;
lateral peridial cells radially elongate; walls of peridial cells
hyaline, less than 1 p thick; uredospores hyaline, very short-
stalked, obovoid, ellipsoid, or subspherical, 14-22 x 23-31 p,
averaging about 17 by 26 p; spore wall colorless, delicately
echinulate, less than 1 p thick.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Dennstaedtia rubiginosa (Kaulf.) Moore, in Colombia,
S. Am. Known only from type locality.
III. Unknown.
Type Locality: Central Andes near Angelopolis, Antioquia, Co¬
lombia, S. Am., on Dennstaedtia rubiginosa. II.
Illustrations: Plate VI, Figs. 25 a-d. — Dietel in Mem. Soc.
Sci. Nat. Neuchatel, 5(2):559. Fig. 66. (1914).
Specimens Examined. —
Colombia: Central Andes near Angelopolis, Antioquia, Aug. 20,
1910, E. Mayor , on Dennstaedtia rubiginosa (II) ; type.
Milesia Dennstaedtiae is inconspicuous in dried plants be¬
cause of the smallness of the uredinia and the long persistence
1932]
FAULL, THE GENUS MILESIA
49
of normal green color in the lesions. The uredinia, however,
are numerous and typical. The cells of their peridia are char¬
acterized by very thin walls.
V. SPECIES ON DRYOPTERIS
- 10. Milesia dilatata Faull, sp. nov. II et III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregati, pustulati, rotundati, 0.1-
0.2 mm. diam., peridio ex cellulis hyalinis irregulariter poly-
gonalibus composito cincti; cellulae peridii 9-14 p diam., pari-
ete circa 1 p crasso. Uredosporae obovatae vel ellipsoideae,
leves, hyalinae, 13-19 x 24-40 p, plus minusve 15 x 30 p, epis-
porio 0.7- 1.0 p crasso. Teleutosporae in hypophyllo intra
cellulas epidermidis evolutae, verticaliter septatae, pluricellu-
lares, leves, hyalinae; celluli teleutosporarum 6-14 x 8-17 p.
Hab. in foliis Dryopteridis spinulosae dilatatae in Oregon,
America bor.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
aggregate on greenish to brown areas of indefinite extent, in¬
conspicuous, pustular, round, 0.1 -0.2 mm. in diameter, covered
by a brownish discolored epidermis which finally ruptures at a
centrally placed stomatic pore; peridium hemispheric, deli¬
cate; peridial cells hyaline, isodiametrically to irregularly
polygonal, 9-14 p across; walls of peridial cells hyaline, thin,
1 p or less; uredospores colorless, white in mass, numerous,
produced singly, short-stalked, obovoid to ellipsoid, 13-19 x
24-40 p, averaging about 15 x 30 p; walls of spores thin, 1 p
or less, smooth.
III. Telia on overwintered fronds, hypophyllous and occa¬
sionally epiphyllous, on indefinite, extensive, brown areas, at
times involving entire pinnae; teliospores within the epidermal
cells, often within the guard cells, hyaline, rounded, or irregu¬
lar in outline and conforming to the shape of the containing
epidermal cell, often completely filling it, one- to many-celled,
with anticlinal septa, a single pore in the outer wall of each
cell; the cells of the teliospores with rather thin, smooth, color¬
less walls, irregularly polygonal except along their free mar¬
gins, 6-14 x 8-17 p.
50
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Hosts and Distribution:
0 and I. Unknown.
II and III. Dryopteris spinulosa (O. F. Muller) Kuntze
var. dilatata (Hoffm.) Underw., in Oregon, U. S.
Type Locality: Zig Zag River, Government Camp, Oregon, U. S.,
on Dryopteris spinulosa var. dilatata. II and III.
Illustrations: Plate III, Figs. 10 a-d; Plate IX, Fig. 35.
Specimens Examined, all on Dryopteris spinulosa var. dilatata. — -
Oregon: Zig Zag River, Government Camp, July 15, 1929, G. D.
Darker (2789), Arnold Arb. Path. Herb. 284 (II, III), telia
on overwintered fronds; type. — Zig Zag River, Government
Camp, July 16, 1929, G. D. Darker (2799), Arnold Arb.
Path. Herb. 294 (II), fronds of current season. — Camp
Creek, Rhododendron, July 20, 1929, G. D. Darker (2839),
Arnold Arb. Path. Herb. 334 (II, III), overwintered fronds
with uredinia and telia, fronds of current season with ure-
dinia. — Camp Creek, Rhododendron, Sept. 6, 1931, J. H.
Fault (II).
Considerable interest attaches to the finding of a new spe¬
cies of Milesia on Dryopteris spinulosa dilatata in America.
Milesia Kriegeriana occurs on Dryopteris spinulosa in Europe,
and more latterly A. Wroblewski, M. Wilson and G. D. Darker
located it on the variety dilatata in Poland, Scotland and Eng¬
land respectively. Two other species, M. intermedia and M.
fructuosa, are described in this paper, the former widely rep¬
resented on D. spinulosa intermedia in northeastern America
and the latter on D. spinulosa fructuosa and D. spinulosa
americana in New England. So now four species of Milesia are
recognized within the spinulosa group of Dryopteris. The
habits and morphological features of these rusts are so different
that specific distinctions are readily drawn. Among the more
important differences the following are noteworthy. The telia
of M. Kriegeriana and likewise also those of M. fructuosa
develop in the fall on fronds of the current season, those of the
others in the spring on overwintered fronds. The uredinia of
M. Kriegeriana are conspicuous, abundant, pustular, many-
spored, and they form at once on new lesions; their uredospores
are constantly spinulose. The uredinia of M. dilatata and of
M. fructuosa are similar, but the uredospores of the former are
perfectly smooth and those of the latter are minutely verrucose
or roughened. The uredinia of M. intermedia are very incon¬
spicuous, very infrequent, submerged and not pustular, few-
1932]
FAULL, THE GENUS MILESIA
51
spored, and they have been seen only on the lesions in over¬
wintered fronds; their uredospores are minutely warted.
Pores in the uredospores of M. dilatata are demonstrable by
boiling the spores in lactic acid. They are faintly visible in
profile after such treatment, as the membranes of the pores
are distended and more hyaline than other areas of the spore
wall.
The teliospores of M. dilatata are usually many-celled.
Single-celled spores are rare; 2 -celled spores are frequent; 6-
to 10-celled spores are abundant. The spores that occur now
and then in the guard cells are usually 2 -celled, and they are
crescentic, conforming perfectly to the containing cells. Few-
celled spores in epidermal cells other than guard cells are
approximately circular in outline in the periclinal plane.
Many-celled spores are sinuously shaped according to the
irregular outlines of the cells in which they are borne.
In passing it should be noted that the D. spinulosa dilatata
referred to above is very close to a fern in Europe known
under this name, but is different from the D. spinulosa dila¬
tata Am. auth. of the region from the Rocky Mountains east¬
ward in North America. The latter has been designated as
Dryopteris spinulosa var. americana (Fisch.) Fernald.
11. Milesia fructuosa Faull, sp. nov. II et III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregati, pustulati, rotundati vel
elongati, 0.15-0.35 mm. diam., peridio ex cellulis hyalinis irreg-
ulariter polygonalibus composito cincti; cellulae peridii 7-12 x
8-15 m, pariete 1.0- 1.3 m crasso. Uredosporae obovatae vel
ellipsoideae, subtiliter verruculosae, hyalinae, 14-21 x 22-36 m,
plus minusve circa 17 x 29 n, episporio circa 1 n crasso. Teleu-
tosporae intra cellulas epidermidis evolutae, amphigenae, prae-
cipue hypophyllae, verticaliter septatae, pluricellulares, leves,
hyalinae; cellulae teleutosporarum 8-16 x 11-27 [ \ .
Hab. in foliis Dryopteridis spinulosae jructuosae in Massa¬
chusetts et Vermont, et in foliis Dryopteridis spinulosae ameri-
canae in Massachusetts et Vermont.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous and often on rhachides and
rhacheolae, subepidermal, scattered or loosely grouped on
52
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
brown or greenish areas of indefinite extent, pustular, round,
0.15-0.35 mm. in width, those on the rhachides and rhacheolae
often linear and up to 3 mm. long, covered by brownish dis¬
colored epidermis which finally ruptures at a centrally placed
stomatic pore or by an irregular slit; peridium firm, hemi¬
spheric; peridial cells hyaline, isodiametrically to irregularly
polygonal, 7-12 x 8-15 p; walls of peridial cells hyaline 1.0- 1.3
p thick; uredospores white, short-stalked, obovoid or ellipsoid,
14-21 x 22-36 p, averaging about 17 x 29 p; walls of spores
about 1 p thick, minutely warted or roughened, more conspic¬
uously so at apical end, exceptionally with occasional, very
short spines.
III. Telia formed in great abundance on fronds of current
season, amphigenous, but mostly hypophyllous, on brown
areas of indefinite extent often involving entire secondary pin¬
nae; teliospores within the epidermal cells, rarely in the guard
cells, hyaline, rounded, or irregular in outline and conforming
to the shape of the containing epidermal cell, usually complete¬
ly filling it, one- to twenty-celled, with vertical septa, a single
pore in the outer wall of each cell; the cells of the teliospores
with thin, smooth, colorless walls, irregularly polygonal except
along their free margins, 8-16 x 11-27 p.
Hosts and Distribution:
O and I. Unknown.
II and III. Dryopteris spinulosa (O. F. Muller) Sw. var.
jructuosa (Gilbert) Trudell, in Massachusetts, Vermont.
II and III. Dryopteris spinulosa var. americana (Fisch.)
Fernald, in Massachusetts, Vermont.
Type Locality: Mt. Greylock, Massachusetts, on Dryopteris
spinulosa var. jructuosa. II and III.
Specimens Examined. —
A. On Dryopteris spinulosa var. jructuosa.
Massachusetts: Mt. Greylock, June 17, 1932, J. H. Fault and
K. S. Chester, on overwintered fronds (II, III); type. — Mt.
Greylock, Aug. 6, 1932, J. H. Fault, on fronds of current
season (II).
Vermont: Searsburg, June 17, 1932, J. H. Fault and K. S. Chester,
on overwintered fronds (II, III). — Mt. Mansfield, July 15,
1932, J. H. Fault and K. S. Chester, on overwintered fronds
(II, III).
B. On Dryopteris spinulosa var. americana.
Massachusetts: Mt. Greylock, July 14, 1932, J. H. Fault and
1932]
FAULL, THE GENUS MILES I A
53
K. S. Chester, on overwintered fronds (II, III). — Mt. Grey-
lock, Aug. 7, 1932, J. H. Fault, on fronds of current season
(II, III).
Vermont: Sherburne Pass, Aug. 10, 1932, on fronds of current
season, /. H. Fault (II, III).
Milesia fructuosa causes large dark blotches on the fronds
that often extend over to and along the rhaohides and rhache-
olae. It is easily distinguished from M. Kriegeriana on Dry-
opteris spinulosa, M. intermedia on D. spinulosa intermedia
and M. dilatata on D. spinulosa dilatata. Its finely roughened
or minutely verrucose uredospores at once set it apart on the
one hand from M. Kriegeriana in which the uredospores are
conspicuously spinulose, and on the other hand from M. dila¬
tata in which the spores are perfectly smooth. Its uredinial
peridium is firmer than in the other species named, and its
teliosporic cells are larger than in M. dilatata. From M. in¬
termedia it differs markedly because of its conspicuous, pustu¬
lar uredinia from which there is a copious discharge of spores,
while in M. intermedia the uredinia are immersed, very incon¬
spicuous, very small, few-spored and rarely seen. Moreover
the teliospores of M. jructuosa are formed on fronds of the
current season, while those of M. intermedia are formed in the
spring on overwintered fronds. The lesions on the new fronds
of the fern host caused by this rust become apparent as early
as midsummer; at once uredinia form and almost simultane¬
ously the epidermal cells begin to fill with teliospores.
Once having recognized M. jructuosa I have found it abund¬
antly represented in the northern Berkshires of Massachusetts
and the Green Mountains of Vermont. At elevations of about
2000 feet and upwards combinations of Balsam Fir with Dry-
opteris spinulosa var. americana or D. spinulosa var. jructuosa
or both are frequent, and in such combinations this rust is com¬
mon on the fern hosts and a white Peridermium equally
common on the Firs. In Sherburne Pass, Vermont, I found
combinations of Balsam Fir with D. spinulosa var. intermedia,
Balsam Fir with D. spinulosa var. americana, and Balsam Fir
with both varieties of ferns. Here there was a great abundance
in 1932 of both Milesia intermedia and M. jructuosa but dis¬
tributed, so far as M. jructuosa was concerned, according to
54
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
its host affinities as recorded in this paper. Further observa¬
tions, including cultural tests, should be made on M. inter¬
media in this connection.
Almost certainly Abies will prove to be the alternate host of
M . jructuosa and it is anticipated that its peridermia occur on
the foliage of the current season only. These conclusions are
based partly on what is known of the life histories of such
rusts as M. intermedia and M. marginalis , and partly on my
finding in the mountains of Massachusetts and Vermont large
quantities of a “white” Peridermium (O and I) on the new
needles of Abies balsamea on trees growing in the midst of
colonies of Dryopteris spinulosa jructuosa and D. spinulosa
americana, respectively, heavily rusted with M. jructuosa and
where no other rusts on ferns were located in the immediate
neighborhoods.
12. Milesia andina Faull, sp. nov. II.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, numerosi, aggregati vel sparsi, pustulati,
rotundati, 0. 1-0.2 mm. diam., peridio ex cellulis hyalinis irreg-
ulariter polygonalibus composito cincti; cellulae peridii 7-15 p
diam., pariete 2-3 p crasso. Uredosporae obovatae, ellip-
soideae vel subglobosae, remote subaculeatae vel verruculosae,
hyalinae, 14-17 x 21-34 p, plus minusve 15 x 26 p, episporio
0. 7-0.8 p crasso. Teleutosporae ignotae.
Hab. in foliis Dryopteridis spec, in Ecuador.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, numerous, ag¬
gregate or scattered on brownish areas of indefinite extent,
often involving entire pinnae, pustular, round, 0.1 -0.2 mm. in
width, covered by a brownish epidermis, usually with centrally
placed stoma tic pore; peridium hemispheric, firm, closely ad¬
herent to epidermis; peridial cells hyaline, isodiametrically to
irregularly polygonal, 7-15 p across; walls of peridial cells
hyaline, 2-3 p thick; uredospores white, short-stalked, obo-
void, ellipsoid or subglobose, 14-17 x 21-34 p, on an average
about 15 x 26 p; walls of spores colorless, thin, 0. 7-0.8 p thick,
sparsely verrucose or subspinulose.
III. Telia unknown.
1932]
FAULL, THE GENUS MILESIA
55
Hosts and Distribution:
O and I. Unknown.
II. Dryopteris spec, (subgenus Lastrea, aff. D. oligocarpa) ,
in Ecuador.
III. Unknown.
Type Locality: Pichincha, Ecuador, on Dryopteris spec. II.
Illustrations: Plate VIII, Figs. 30 a-d.
Specimens Examined. —
Ecuador: Pichincha, G. Lager keim, on Dryopteris spec. (II); type.
— Quito, June 1890, G. Lager keim (II), Herb. Barbey-
Boissier, Univ. of Geneva.
A part of Lagerheim’s collection was deposited in the Her¬
barium of the New York Botanical Garden, a portion of which
was shared with the Arthur Herbarium. The original packet
bore the name Uredo Scolopendrii (Fuckel). Professor
Arthur informs me that he submitted the material to Professor
L. M. Underwood, who determined the host as a species of
“Phegopteris” ( Dryopteris ) .
13. Milesia carpatica (Wroblewski) Faull, n. comb. II and
III.
Milesina carpatica A. Wroblewski in Spraw. Kom. Fizyor.
Akad. Umiejetnosci Krakowie, 47:166, 178 (1913). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, pustular, punc¬
tate, round, 0.08 to 0.2 mm. in diameter, covered by brownish
colored epidermis with centrally placed stomatic pore, often
occurring singly at the center of small, sharply defined, dark
brown spots, but also loosely aggregate on brownish areas of
indefinite extent that appear to result from the extension and
coalescence of the primary lesions; peridium hemispheric, deli¬
cate; peridial cells hyaline, isodiametrically to irregularly poly¬
gonal, 6 to 12 p across; walls of peridial cells hyaline, 0.5 to
1.0 p thick; uredospores colorless, very thin-walled (0.5-0. 7
p), short-stalked, obovoid, ellipsoid or subglobular, 11-17 x
14-27 p, averaging about 14 x 20 p; walls of spores with short,
delicate echinulations.
III. Telia amphigenous, mostly hypophyllous, on overwin¬
tered fronds, on indefinite, extensive brown areas; teliospores
within the epidermal cells, exceptionally in guard cells, hya¬
line, rounded, or irregular in outline and conforming to the
shape of the containing epidermal cell, often completely filling
56
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
it, one- to many-celled (up to 60 or more), with anticlinal
septa; the cells of the teliospores with thin, smooth, colorless
walls, irregularly polygonal except along their free margins,
5.5-11 x 8-15 p.
Teleutosporae intra cellulas epidermidis evolutae, amphig-
enae, praecipue hypophyllae, verticaliter septatae, pluricellu-
lares, leves, hyalinae; cellulae teleutosporarum 5.5-11 x 8-15 p*
Hab in foliis Dryopteridis Filicis maris in Czechoslovakia.
Hosts and Distribution:
O and I. Unknown.
II. Dryopteris Filix mas (L.) Schott, in Poland, Czecho¬
slovakia.
III. Dryopteris Filix mas (L.) Schott in Czechoslovakia.
Type Locality: Near Kolomea, Poland, on Dryopteris Filix mas.
II.
Illustrations: Plate II, Figs. 8 a-d. — Wroblewski in Bull. Acad.
Sci. Cracovie, Oct.-Dec. 1915. PI. 8, Fig. 2 A (1916).
Exsiccati: Syd. Ured. 2839.
Specimens Examined, all on Dryopteris Filix mas. —
Czechoslovakia: Winterberg, July 2, 1932, G. D. Darker, on over¬
wintered fronds (II, III); type for teliospores. — Kralovany,
Slovakia, July 27, 1932, G. D. Darker (II).
Poland: Kniazdwor, Kolomea, May 1912, A. Wroblewski (II);
part of type. — Pohulanka, Lwow, Sept. 3, 1916, A. Wroblew¬
ski (II). — Kniazdwor, Kolomea, Sept. 1912, A. Wroblewski
(II). — Kniazdwor, Kolomea, Sept. 1913, A. Wroblewski, Syd.
Ured. 2839 (II). '
Milesia carpatica is readily distinguished from M. Kriegeri-
ana, which also occurs on Dryopteris Filix mas, by its con¬
stantly much smaller, more delicately echinulate uredospores,
and by its much smaller-celled teliospores. Whether or not the
teliospores of M. carpatica are formed in the fall on fronds of
the current season as for M. Kriegeriana I am unable to state.
The teliosporic material available for my study was on over¬
wintered fronds, but it was collected so late in the season that
no conclusions could be drawn as to when the teliospores
might have first appeared. No teliospores, however, were
found on fronds of the current season collected in the fall, at
dates no later than those at which the teliospores of M. Krie¬
geriana appear. Wroblewski (50) refers to teliospores in M.
carpatica, but his description and drawings indicate that what
he saw were bodies quite apart from the rust.
1932]
FAULL, THE GENUS MILESIA
57
Incidentally it may be observed that in the two collections of
the five examined by me (collections made by Dr. G. D.
Darker) in which the fern sori were present the indusia were
glandular. I am informed by Mr. C. A. Weatherby of the
Gray Herbarium that Dryopteris Filix mas in Europe some¬
times carries glandular indusia. One is led to wonder whether
or not such occurrences are peculiar to a form or variety of
D. Filix mas. In any case it would be interesting to know
whether or not M. carpatica is restricted to plants of D. Filix
mas with glandular indusia and M. Kriegeriana to plants with
smooth indusia.
I am under obligations to Director Wroblewski for ample
collections of uredosporic material from Poland and to Dr.
G. D. Darker for teliosporic material from Czechoslovakia.
14. Milesia carpatica (Wrob.) Faull var. erythrosora Faull,
var. nov. II et III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregate minuti, pustulati, rotun-
dati, 0.1 -0.2 mm. diam., peridio ex cellulis hyalinis irregulariter
polygonalibus composito cincti; cellulae peridii 6-12 |j diam.,
pariete 0.5-0. 7 [i crasso. Uredosporae obovatae, ellipsoideae vel
rarius subglobosae, remote et subtilissime echinulatae, hya-
linae, 14-20 x 21-28 m, plus minusve 16 x 25 m, episporio 0.7-
0.9 n crasso. Teleutosporae in hypophyllo intra cellulas epi-
dermidis evolutae, verticaliter septatae, 1 — 50-cellularibus,
leves, hyalinae; cellulae teleutosporarum 6-11 x 7-14 p.
Hab. in foliis Dryopteridis erythrosorae in Japonia.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, punctate, round,
0.1 -0.2 mm. in diameter, covered by brownish covered epider¬
mis with centrally placed stomatic pore, scattered or aggregate
on brownish areas of indefinite extent; peridium hemispheric,
delicate; peridial cells hyaline, isodiametrically to irregularly
polygonal, 6 to 12 m across; walls of peridial cells hyaline, less
than 1 m thick; uredospores colorless, very thin-walled (less
than 1 m thick), short-stalked, obovoid, ellipsoid, or rarely
subglobular, 14-20 x 21-28 |j, averaging about 16 x 25 m; walls
of spores with short, scattered, delicate echinulations.
III. Telia hypophyllous, on overwintered fronds, on in-
58
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
definite, extensive brown areas; teliospores within the epider¬
mal cells, frequently in the guard cells, hyaline, rounded, or
irregular in outline and conforming to the shape of the con¬
taining epidermal cell, often completely filling it, few- to many-
celled (up to 50 or more), with anticlinal septa; the cells of
the teliospores with thin, smooth, colorless walls, irregularly
polygonal except along their free margins, 6-11 x 7-14 [i.
Hosts and Distribution:
O and I. Unknown.
II and III. Dryopteris erythrosora (Eaton) Kuntze, in
Japan.
Type Locality: Tottori, Japan, on Dryopteris erythrosora.
Specimens Examined. —
Japan: Tottori, March 21, 1930, Naohide Hiratsuka, on Dryopteris
erythrosora (II, III), on overwintered fronds; part of type.
The cells of the teliospores of Milesia carpatica var. erythro¬
sora are exceptionally small, constituting one of the few in¬
stances for the genus Milesia in which the teliospores are of
some value in the diagnosis of species.
Regarding the name to be given to this rust it is plainly not
M . Kriegeriana, which it recalls in some respects. But its ure-
dinia, uredospores, uredospore markings and telial cells are
distinctly smaller than those of M. Kriegeriana. If its telia
are formed, not in the fall as is true of the latter species, but
in the spring on overwintered fronds, that would constitute an¬
other distinction. Its teliospores, uredinia and uredo¬
spores resemble more closely those of M. carpatica. The main
differences comprise its somewhat larger uredospores, and the
rather dissimilar lesions it causes. These are perhaps too
slight to warrant complete specific separation, but sufficiently
significant to warrant varietal recognition. In any case it is
reasonable to anticipate from our experiences with cross cul¬
tures of species of Milesia, that the two are not interchangeable
as to hosts. All in all it seems sound to tentatively designate
this rust on Dryopteris erythrosora a variety of M. carpatica.
15. Milesia Kriegeriana (Magn.) Arthur, excl. descript., in
Mycologia, 7:176 (1915). (II). I, II and III.
Melampsorella Kriegeriana Magnus in Ber. Deutsch. Bot.
Ges. 19:581 (1901). II and III.
1932]
FAULL, THE GENUS MILESIA
59
Hyalopsora Kriegeriana (Magnus) Fischer in Die Ure-
dineen der Schweiz, p. 538 (1904). II and III.
Milesina Kriegeriana Magnus in Ber. Deutsch. Bot. Ges.
27:325 (1909). II and III.
I. Aecia demonstrated by Dr. Eugene Mayor in 1931. De¬
scription not yet available. Spermogonia are said to be
lacking.
II. Uredinia hypophyllous, subepidermal, numerous, scat¬
tered or loosely grouped on greenish to brown areas of in¬
definite extent, pustular, round, 0.1 -0.3 mm. in width, covered
by brownish discolored epidermis with centrally placed sto-
matic pore; peridium hemispheric, delicate; peridial cells, hya¬
line, isodiametrically to irregularly polygonal, overlapping, 7
to 14 p across, lateral ones radially elongated; walls of peridial
cells hyaline, about 1 p thick; uredospores colorless, produced
singly, with stalks 2 to 8 p long, obovoid to ellipsoid, 15-22 x
23-48 p, on an average about 18 x 33 p; walls of spores thin,
1 p or less, echinulate.
III. Telia formed in fall on fronds of current season, hypo-
phyllous, on brown areas of indefinite extent; teliospores with¬
in the epidermal cells, sometimes within even the guard cells,
hyaline, rounded, or irregular in outline and conforming to the
shape of the containing epidermal cell, often completely filling
it, one- to forty-celled, with anticlinal septa; the cells of the
teliospores with thin, smooth, colorless walls, irregularly polyg¬
onal except along their free margins, 6-16 x 8-20 p.
Hosts and Distribution:
I. Abies alba Mill., in Switzerland.
II. Dryopteris spinulosa (O. F. Muller) Kuntze, in Ger¬
many, Austria, Switzerland, Denmark, Finland, Poland, Eng¬
land, France.
Dryopteris spinulosa var. dilatata (Hoffm.) Underw. in
Poland, Scotland, England.
Dryopteris Filix mas (L.) Schott, in Poland, Switzerland,
England, France.
III. Dryopteris spinulosa (O. F. Muller) Kuntze, in Ger¬
many, Switzerland, Poland, England, France.
Dryopteris spinulosa var. dilatata (Hoffm.) Underw., in
Poland.
Dryopteris Filix mas (L.) Schott, in Poland, Switzerland,
France.
60
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Type Locality: Konigstein, Germany, on Dryopteris spinulosa.
II and III.
Illustrations: Plate III, Figs. 9 a-d. — P. Magnus in Ber.
Deutsch. Bot. Ges. 19: PI. 33 (1901. — P. and H. Sydow in Monog.
Ured. 3, PI. 21, Fig. 164, p. 475 (1915).
Exsiccati: (a) On Dryopteris spinulosa : Krieg. Fg. Saxon. 856,
under name Uredo Scolopendrii (Fuckel) Schroet., 1711, — Rabh.
Fg. Eur. 4242, under name Uredo Scolopendrii (Fuckel) Schroet. —
Syd. Ured. 1634, under name Melampsorella Kriegeriana Magn.
(b) On Dryopteris Filix mas : Syd. Ured. 2598, 2642. — Vestergren
Microm. Rar. Sel. 1704. — Syd. Myc. Germ. 1329.
Specimens Examined. —
A. On Dryopteris spinulosa.
Germany: Lattengrund, Saxony, Oct. 12, 1892, and Sept. 16, 1893,
W. Krieger, Krieg. Fg. Saxon. 856 (II, III). — Saxony, Sept.
1898, W. Krieger, Rabh. Fg. Eur. 4242 (II). — Polenzthal
near Konigstein, Sept. 1901, W. Krieger, Syd. Ured. 1634
(II, III); type locality. — Uttewalder Grunde, Saxony, Oct.
and Nov. 1901, W. Krieger, Krieg. Fg. Saxon. 1711 (II, III).
Poland: Kniazdwor, near Kolomea, Sept. 1913, A. Wroblewski (II,
III). — Kielce, Sept. 29, 1925, Wanda Konopacka, Herb. W.
Siemaszko (II).
Switzerland: Canton of Neuchatel, Aug. 1, 1909, E. Mayor (II). —
Bern, June 2, 1932, G. D. Darker (II, III).
England: Cold Harbour, Surrey, Nov. 8, 1931, G. D. Darker (II).
— Cold Harbour, Surrey, Nov. 12, 1931, G. D. Darker (II).
— Brockenhurst, Hants Co., Nov. 25, 1931, G. D. Darker
(II, III). — Lytchett Minster, Dorset Co., Nov. 26, 1931,
G. D. Darker (II).
France: Puivert, Aude, May 4, 1932, G. D. Darker (II). — Axat,
Aude, Apr. 30, 1932, G. D. Darker (II, III).
B. On Dryopteris spinulosa var. dilatata.
Poland: Zawojela in eastern Carpathians, undated, A. Wroblewski
(II, III).
Scotland: Kelso, Aug. 19, 1925, M. Wilson, Kew Herb, and Herb.
Bur. PI. Ind., U. S. Dept, of Agr., Washington, D. C. (II).
England: Lytchett Minster, Dorset Co., Nov. 26, 1931, G. D.
Darker (II).
C. On Dryopteris Filix mas.
Poland: Oseredok, near Kolomea, Sept. 1913, A. Wroblewski (II,
III). — Kniazdwor, near Kolomea, Sept. 1913, A. Wroblew¬
ski (II). — Kniazdwor, Kolomea, Sept. 1913, A. Wroblewski ,
Syd. Ured. 2642 (II). — Skomorochy, Sokal, Sept. 28, 1913,
A. Wroblewski (II). — Olszanica, near Lesko, Oct. 18, 1917,
A. Wroblewski (II, III).
Switzerland: Creux du Vau, Neuchatel, Oct. 12, 1913, E. Mayor,
Syd. Ured. 2598 (II, III). — Perreux sur Boudry, Neuchatel,
1932]
FAULL, THE GENUS MILES1A
61
Sept. 15, 1917, E. Mayor (II, III). — Creux du Vau, Neu-
chatel, Oct. 19, 1913, E. Mayor, Vestergren Microm. Rar.
Sel. 1704 (II, III).
France: Ltitzelhausen, Kreis Molsheim, Alsace, Dec. 7, 1914, A.
Ludwig, Syd. Myc. Germ. 1329 (II, III). — Ax-les-Thermes,
Ariege, May 6, 1932, G. D. Darker (II, III). — Axat, Aude,
May 1, 1932, G. D. Darker (II, III).
England: Tintern, April 1925, E. M. Wakefield, Herb. Bot. Reg.
Kew (II). — Brockenhurst, Hants Co., Nov. 25, 1931, G. D.
Darker (II). — Lytchett Minster, Dorset Co., Nov. 26, 1931,
G. D. Darker (II).
MUesia Kriegeriana was described under the name Melamp-
sorella Kriegeriana by P. Magnus (35) from uredinial and
telial material on Dryopteris spinulosa sent from Saxony by
W. Krieger. Teliospores were observed the first time for any
species of Milesia, and figured from material collected in No¬
vember. Magnus also germinated the teliospores and found
that they produced ordinary basidia, one from each cell. This
rust, along with M. Feurichii, was later included, as type, by
Magnus (37) in his new genus Milesina.
One interesting feature about M. Kriegeriana is the fact
that its teliospores develop in the fall on the fronds of the cur¬
rent season. I have seen telial material from Germany,
France, Switzerland, Poland and England, and in each instance
it was collected on fronds of the current season in the late fall.
As M. Kriegeriana was the first species of the genus in which
teliospores were found, this occurrence led to the erroneous
belief that such is the typical habit of the genus. As a matter
of fact that is not the case, for with few known exceptions
they form in the following spring on affected, overwintered
fronds. As to the viability of the teliospores Magnus (35)
claims to have been able to secure immediate germination in
the fall, but he did not indicate to just what extent. Dietel
(11) states that he found the teliospores of M. Kriegeriana to
be still ungerminated in the early spring on overwintered
fronds, a statement confirmed by my own observations, and
concludes that for the germination of a part of them at least
a winter resting period is the rule in nature. Regardless of the
soundness of Dietel’s deduction it is true that germination pre¬
vious to the unfolding of the needles of Abies in the spring
would be a futile habit.
62
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
In 1913 Wroblewski collected a Milesia on Dryopteris Filix
mas in Poland, and it was referred by Sydow (44) to Milesina
Kriegeriana. The same species has subsequently been found
in France, Switzerland and England, and collected further in
Poland. After carefully examining several collections from
these countries I have reached the conclusion that Sydow’s de¬
termination (but under name Milesia Kriegeriana) should
stand, awaiting the completion of life history studies. Ure-
dinia, uredospores, and telia, both as to morphology and time
of development, answer closely to similar features in M. Krie¬
geriana.
Wroblewski likewise discovered a Milesia on Dryopteris
spinulosa var. dilatata in Poland. Through his courtesy I have
had an opportunity of studying a part of his collection. This
material contains both uredinia and telia on what appear to be
fronds of the current season. I have also examined two col¬
lections from Great Britain, one made by Dr. Malcolm Wilson
from the extreme south of Scotland, and the other by Dr.
G. D. Darker from Dorset Co., England. The resemblance to
M. Kriegeriana is close, and I would hesitate to do other than
let this rust tentatively pass as M. Kriegeriana.
A Milesia on Dryopteris spinulosa var. dilatata from the
State of Oregon was collected several times by Dr. Darker in
the summer of 1929, and again by myself in 1931. This, how¬
ever, is quite distinct from M. Kriegeriatna and is described in
the present paper under the name M. dilatata, n. sp.
So far as is known M. Kriegeriana has not been found else¬
where than in Europe. Arthur (2, 5) identified a Milesia
on Dryopteris marginalis in America as M. Kriegeriana, but
that is an entirely different species. It is recorded in this
paper as M. marginalis, n. sp.
I have likewise found a Milesia many times, often in great
abundance, on Dryopteris spinulosa var. intermedia in the
eastern parts of Canada and the United States. It, too, has
passed current under the name M. Kriegeriana. But with
fuller knowledge of this rust and of the real M. Kriegeriana I
have unhesitatingly concluded that it is neither M. marginalis
nor M. Kriegeriana. It is described in this paper as M. inter¬
media, n. sp.
The aecial host of M. Kriegeriana was experimentally dem-
1932]
FAULL, THE GENUS MILESIA
63
onstrated by Dr. Eugene Mayor in 1931 according to informa¬
tion conveyed in a letter from him. He writes that by sowing
spores from white peridermia of the Peridermium pseudocol-
umnare type growing on Abies alba, on to fronds of Dryop-
teris spinulosa, lesions developed within a brief interval, and
that these lesions bore typical uredinia of M. Kriegeriana .
This result is quite in keeping with what has been anticipated
regarding the alternate host of this well-known fern rust. One
interesting observation made by him is to the effect that the
peridermia from which he cultured were not accompanied by
spermogonia. We shall await with interest his cultures made
in the reverse direction.
16. Milesia consimilis Arthur in Mycologia, 7:176 (1915).
II.
Milesina consimilis (Arthur) Trotter in Saccardo’s Syll.
Fung. 23:846 (1925).
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely aggregate on discolored areas of indefinite extent, pus¬
tular, round to slightly elongate, 0.2-0. 3 mm. in width, cov¬
ered by yellowish epidermis, which ruptures irregularly; perid-
ium hemispheric, colorless, firm; peridial cells isodiametrically
to irregularly polygonal, 8 to 16 n across, with walls up to 2 p
thick; uredospores obovoid, or more rarely ellipsoid or sub-
globular, white, 17-23 x 23-34 p , averaging about 19 x 28 p;
walls of spores hyaline, about 1.5 p thick, strongly and rather
sparsely to profusely echinulate.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Dryopteris patens (Sw.) Kuntze, in Jamaica.
III. Unknown.
Type Locality: Whitfield Hall, Jamaica, on Dryopteris patens.
II.
Illustrations: Plate VI, Figs. 21 a-d.
Specimens Examined, all on Dryopteris patens. —
Jamaica: Whitfield Hall, April 22, 1903, L. M. Underwood (II);
part of type. — Cinchona, alt. 5000 feet, Feb. 20, 1900, Wil¬
lard N . Clute, Herb. N. Y. Bot. Gard. 364 (II).
64
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Through the courtesy of Professor J. C. Arthur and of Dr.
F. J. Seaver I have had the opportunity of examining the
original material collected by L. M. Underwood on Dryopteris
patens in Jamaica. My description corresponds closely with
that given by Professor Arthur (2). The main departure is
with respect to the peridium. This I find to be relatively firm
and thick-walled. It quite readily separates from the overly¬
ing epidermis. It may be noted that the walls plainly show
their double composition, which is in keeping with the fact
that the partition separating adjacent peridial cells consists of
the walls of two contiguous cells of independent origin.
Milesia consimilis is so far known from Jamaica only, and
on the one host Dryopteris patens. Arthur (2, p. 325) at one
time recognized a wider distribution both as to hosts and lo¬
calities, but later (5, p. 687) appears to have rightly revised
his earlier opinion. It should be noted that in 1910 Christen¬
sen divided the widely distributed D. patens into two species,
D. patens (Sw.) Kuntze and D. normalis Christ., both re¬
corded in the fern flora of Jamaica as well as elsewhere; but
they are not separable in such fragments as are found in rust
herbaria.
17. Milesia intermedia Faull, sp. nov. O, I, II et III.
Peridermium balsameum Peck in Rep. N. Y. State Mus.
27:104 (1875), in part.
Milesia Kriegeriana (Magnus) Arthur in N. Am. Flora,
7:686 (1925), as to description in part. O, I, II and III.
Pycnidia hypophylla, numerosa, inconspicua, hyalina, hemi-
sphaerica, subcuticularia, 84-137 m lata x 59-84 p alta, plus
minusve 110 x 71 p; aecidia hypophylla, maeulis flavidis insi-
dentia, secus series duas irregulares disposita, cylindracea
vel parum lateraliter compressa, 0.3-0. 4 mm. diam. x 0. 2-1.0
mm. alta, alba, ad apicem dehiscentia; cellulae peridii sub-
imbricatim positae, 19-23 x 25-57 p, pariete exteriore levi 2-3 p
crasso, interiore 4. 5-5. 5 p crasso papillis varie elongatis dense
rugosis. Aecidiosporae globosae, ovatae vel ellipsoideae, dense
verrucosae, hyalinae, 19-23 x 21-31 p, plus minusve 21 x 27 p,
episporio 1.0- 1.5 p crasso. Sori uredosporiferi parvi ideoque
difficile visi, minuti, hypophylli, epidermide tecti, sparsi, im-
mersi et non pustulati, rotundati, 0.08-0.12 mm. diam., peridio
ex cellulis hyalinis irregulariter polygonalibus composito cine-
1932]
FAULL, THE GENUS MILESIA
65
ti; cellulae peridii non imbricatae, 7-14 p diam., pariete 0.5-
0.8 p crasso. Uredosporae irregulariter obovatae vel ellip-
soideae, remote et subtilissime verruculosae, hyalinae, 14-22 x
20-34 p, plus minusve 16 x 28 p, episporio 0.6-0.9 p crasso.
Teleutosporae intra cellulas epidermidis evolutae, hypophyllae,
verticaliter septatae, pluricellulares, leves, hyalinae; cellulae
teleutosporarum 9-16 x 12-22 p.
Hab. in foliis Abietis balsameae et Dryopteridis spinulosae
intermediae in Ontario, Quebec, Nova Scotia, Maine, New
Hampshire, Vermont, Massachusetts et New York.
O. Spermogonia on needles of current season, hypophyl-
lous, very abundant, inconspicuous, colorless, plane, outer
zone hyaline or watery-looking in fresh condition, immersed,
inverted hemispherical in sectional view, subcuticular, 84-137 p
broad by 59-84 p deep, averaging 1 10 p broad by 71 p deep;
apical pore slit-like, 3. 3 -8. 4 x 15-60 p, parallel to long axis of
needle; spermatiophores unbranched, septate, producing the
spermatia catenulately ; spermatia hyaline, unicellular, nar¬
rowly elliptical, 1.5-1. 6 x 4. 5-7.0 p.
I. Aecia hypophyllous on needles of current season, in two
rows, on yellowish discolored portions of affected needles,
white, circular or slightly compressed laterally in transverse
section, cylindrical, 0.3 to 0.4 mm. in diameter by 0.2 to 1.0
mm. high; peridium colorless, delicate, rupturing at the apex;
peridial cells polygonal, elongated vertically, overlapping, in a
single layer, 19-23 x 25-57 p, with outer walls smooth, 2-3 p
thick, with inner walls closely marked by tubercles and very
short irregularly oriented ridges, 4. 5-5. 5 p thick; aeciospores
globose, ovoid or ellipsoid, colorless, 19-23 x 21-31 p, mostly
somewhat elongated and about 21 x 27 p, finely and closely
verrucose, with the tubercles somewhat deciduous, one side
partly smooth; wall colorless, thin, 1.0-1. 5 p thick including
tubercles.
II. Uredinia infrequent, not easily located, hypophyllous,
subepidermal, scattered on the greenish to brownish lesions,
immersed, not pustular, round, punctate, 0.08-0.12 mm. in
diameter, covered by epidermis with centrally placed stomatic
pore and concolorous with contiguous epidermis; peridium al¬
most plane, not hemispheric, delicate; peridial cells hyaline,
isodiametrically to irregularly polygonal throughout, not over-
66
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
lapping, 7 to 14 p across; walls of peridial cells hyaline, less
than 1 m thick; uredospores colorless, extruded in delicate
tendrils, white in mass, short-stalked, irregularly obovoid or
ellipsoid, 14-22 x 20-34 p, averaging about 16 x 28 p; walls of
spores thin, less than 1 p thick, almost smooth, very minutely
and sparsely warted.
III. Telia hypophyllous, abundant on overwintered fronds,
on dark brown, clearly defined, large spots, often involving
entire secondary pinnae; teliospores within the epidermal cells,
hyaline, rounded, or irregular in outline and conforming to the
shape of the containing epidermal cell, at times completely fill¬
ing it, one- to many-celled, with vertical septa, a single pore in
the outer wall of each cell; the cells of the teliospores with
thin, smooth, colorless walls, irregularly polygonal except
along their free margins, 9-16 x 12-22 p.
Hosts and Distribution:
O and I. Abies balsamea (L.) Mill., in Ontario, Quebec,
Nova Scotia, Maine, New Hampshire, New York.
II and III. Dryopteris spinulosa (O. F. Muller) Sw. var.
intermedia (Muhl.) Underw., in Ontario, Quebec, Nova Scotia,
Maine, New Hampshire, Vermont, Massachusetts.
Type Locality: Bear Island, L. Timagami, Ontario, Canada, on
Dryopteris spinulosa var. intermedia. O, I, II and III.
Illustrations: Plate III, Figs. 11 a-d. — L. M. Hunter in Bot.
Gaz. 83: Fig. 9 (1927).
Specimens Examined. —
O and I. On Abies balsamea.
Ontario: L. Timagami, various dates from July 29 to Sept. 12,
1924-1928, /. H. Fault, Herb. J. H. Faull 4933, 4941, 4953,
4955, 7198, 7199, 7266, 7268, 7300, 7393, 8051.— Cultures,
Aug. 1924, J. H. Faull and W. R. Watson, Herb. J. H. Faull
7312-7320.
Quebec: Shickshocks, Gaspe, Sept. 8, 1928, J. H. Fatdl, Herb.
J. H. Faull 8647a.
Nova Scotia: Liscomb Park, Guysboro Co., July 30, 1929, /. H.
Faull, Herb. J. H. Faull 9272.
Maine: Oquossoc, crop of 1930 (old), Apr. 26, 1931, J. H. Faull,
Herb. J. H. Faull 9830.
New Hampshire: Whitefield, Aug. 25, 1925, /. H. Faull, Herb.
J. H. Faull 7377, 7379.
New York: Adirondack Mts., 1873 (?), C. Peck, Herb. N. Y. State,
and identified by Miss L. M. Hunter.
II. On overwintered fronds of Dryopteris spinulosa inter¬
media.
1932]
FAULL, THE GENUS MILESIA
67
Ontario: L. Timagami, June 14, 1926, J. H. Fanil, Herb. J. H.
Faull 7527.
New Hampshire: Tuckerman Ravine Trail, Mt. Washington, July
6, 1931, /. H. Faull, Herb. J. H. Faull 9933, 9924.
Vermont: Mt. Mansfield, July 15, 1932, /. H. Faull and K. S.
Chester.
Massachusetts: Becket, July 12, 1932, /. H. Faull and K. S.
Chester.
III. On overwintered fronds of Dryopteris spinulosa inter¬
media.
Ontario: L. Timagami, various dates, June 1924 to 1927, J. H.
Faull, Herb. J. H. Faull 4871, 7371 (type), 7373 (type),
7527. — Cultures, June 1925, J. H. Faull and W. R. Watson,
Herb. J. H. Faull 8260, 8267, 8270, 8272. — Cultures, June
1928, E. H. Bensley, Herb. J. H. Faull 9850 c, d.
New Hampshire: Tuckerman Ravine Trail, Mt. Washington, July
6, 1931, J. H. Faull, Herb. J. H. Faull 9933.
Vermont: Mt. Mansfield, July 15, 1932, /. H. Faull and K. S.
Chester. — Sherburne Pass, Aug. 11, 1932, /. H. Faull.
Massachusetts: Becket, July 12, 1932, J. H. Faull and K. S.
Chester.
Milesia intermedia is, perhaps, the most frequent white-
spored rust to be found throughout the range of Abies bal-
samea, and in some areas it is very abundant. Yet, it has not
been recognized because the white-spored peridermia on Abies
balsamea, so long designated Peridermium balsameum Peck,
have been passed by as some species of Uredinopsis, and the
fern host has not been collected because of lack of visible rust
fructifications. My investigations of this rust began with a
search for telia on overwintered fronds, and then having found
them inoculations were made on Abies. A study of the organs
that developed on the needles, revealed the characters by which
this rust can be distinguished from other white-spored rusts on
the Fir. These are comprised in the large numbers of spermo-
gonia on a needle, their size, distribution and structure.
Milesia intermedia on Dryopteris spinulosa intermedia and
M. marginalis on D. marginalis in America are perfectly dis¬
tinct from each other and respectively from the European M.
Kriegeriana; but because of imperfect acquaintance with them
heretofore they have been erroneously referred to the last
named species. In order to give a clear picture of the three
species their distinctive characters are presented side by side
in tabular form as follows:
68
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
TABLE 5
M. intermedia M. marginaHs M. Kriegeriana
Spermogonia
Location
Size — range
Size — average
Frequency
Uredinia
Form
Size
Frequency
Conspicuousness
Occurrence
Upper surface
Peridium
Uredospores
Size — range
Size — average
Markings
Numbers
T elios pores
Occurrence
Incubation period
on Abies
Hypophyllous
84-137 broad x
59-84 p high
110 x 71 p
Crowded
Vesicular (i.e. im¬
mersed) — no beak
0.08-0.12 mm.
Rare
Concealed, evident
only from extruded
spores
Overwintered
fronds
Plane
Cells shallow and
not overlapped
14-22 x 20-34 p
16 x 28 p
Punctate warts
Scanty
Overwintered
fronds
33-36 days
Amphigenous
129-168 x 92-
134 p
147 x 106 p
Frequent, but
much fewer
Vesicular to slight¬
ly pustular — sem¬
blance of beak
0.09-0.25 mm.
Abundant
Inconspicuous
Fronds of current
season (?) Abun¬
dant on over¬
wintered fronds
Slightly convex
Cells deep and
strongly overlapped
15-27 x 27-51 p
19 x 36 p
Low spines and
spiny warts
Rather few
Overwintered
fronds
41-51 days
None
Pustular
0. 1-0.3 mm.
Abundant
Conspicuous
Abundant on fronds
of current season.
Also on over¬
wintered fronds
Hemispheric
Cells somewhat
overlapped
15-22 x 23-48 p
18 x 33 p
Strongly spinulose
Very numerous
Fronds of current
season
Not known
Two of the most remarkable features of M. intermedia are
the paucity and the inconspicuousness of the uredinia. They
are apparently rare, and so reduced in size as to give the im¬
pression that they are all but eliminated from this species.
So far I have located them on overwintered fronds only, and
have been able to do so only by looking for extruded spore
masses; in addition to their minuteness they are submerged,
causing no external or pustular swelling, an almost exceptional
feature for the genus. The spore masses can be detected only
with a hand lens, for the number of spores produced by a
single uredinium is exceedingly small.
The teliospores on the other hand are very abundant in large
brown lesions on affected fronds. The spotting of the fronds
1932]
FAULL, THE GENUS MILESIA
69
occurs in the fall on fronds of the current season, but the telio-
spores do not develop until the following spring — at just about
the time that needles of the Fir are unfolding. Incidentally it
may be noted that the peridermia on Abies are rather late in
appearing, usually not before August or the latter half of July,
and collections have been made as late as September.
There are indications that Dryopteris spinulosa americana
may carry this rust, but the evidence is not yet complete.
18. Milesia marginalis Faull & Watson, sp. nov. O, I, II
et III.
Milesina marginalis Faull & Watson, inedit., in Faull, J. H.,
Proc. Internat. Congr. Plant Sci. (1926) 2:1735 (1929).
Milesia Kriegeriana Arthur in Mycologia, 7:176 (1915),
not Magnus. II.
Milesia Kriegeriana Arthur in N. Am. Flora, 7:686 (1925),
as to description, in part. O, I, II and III.
Pycnidia amphigena, praecipue hypophylla, inconspicua,
hyalina, immersa, hemisphaerica, 125-168 p lata x 90-134 p
alta, plus minusve 147 x 106 p; aecidia hypophylla, maculis
flavidis insidentia, secus series duas disposita, cylindracea vel
parum lateraliter compressa, 0.3 -0.4 mm. diam. x 0.4- 1.0 mm.
alta, alba, ad apicem dehiscentia; cellulae peridii subimbri-
catim positae, 20-40 x 30-50 p, pariete exteriore levi 1. 5-2.0 p
crasso, interiore 4-5 p crasso papillis varie elongatis dense
rugosis. Aecidiosporae globosae, ovatae vel ellipsoideae, dense
et subtiliter verrucosae, hyalinae, 21-28 x 21-34 p, praecipue
subglobosae et plus minusve 24 x 28 p, episporio sine papillis
circa 1 p crasso. Sori uredosporiferi numerosi, inconspicui,
hypophylli, epidermide tecti, sparsi vel aggregati, immersi vel
leviter pustulati, globoso-conici, 90-250 p lati x 100-125 p alti,
peridio ex cellulis hyalinis, peridio inferne ex cellulis radiatim
elongatis superne irregulariter polygonalibus composito cincti;
cellulae peridii forte imbricatae, 6-10 p diam., pariete 1 p vel
minus crasso. Uredosporae irregulariter obovatae vel ellipsoi¬
deae, remote echinulatae vel crasse aculeatae, hyalinae, 15-27 x
27-51 p, plus minusve 19 x 36 p, episporio 0. 7-0.9 p crasso.
Teleutosporae intra cellulas epidermidis evolutae, amphigenae,
praecipue hypophyllae, verticaliter septatae, pluricellulares,
leves, hyalinae; cellulae teleutosporarum 8-16 x 12-25 p.
Hab. in foliis Abietis balsameae et Dryopteridis marginalis
in Ontario, Quebec, Massachusetts, New Hampshire et New
York.
70
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
O. Spermogonia on needles of current season, amphigenous
but mostly hypophyllous, numerous, inconspicuous, colorless,
outer zone hyaline or watery-looking in fresh condition, im¬
mersed, inverted hemispherical in sectional view, covered by
the cuticle and the intermediate layer of external epidermal
wall, 125 to 168 p broad by 90-134 p deep, averaging 147 p
broad x 106 p deep; apical pore slit-like, 1. 7-5.0 x 20-50 p,
parallel to long axis of needle; spermatiophores unbranched,
septate, producing the spermatia catenulately; spermatia hya¬
line, unicellular, narrowly ellipsoid, 1.6 x 5. 0-9.0 p.
I. Aecia hypophyllous, on needles of current season, in
two rows, on yellowish discolored portions of affected needles,
white, circular or somewhat laterally compressed in transverse
section, cylindrical, 0.3 to 0.4 mm. in diameter by 0.4 to 1.0
mm. high; peridium colorless, delicate, rupturing at the apex;
peridial cells polygonal, elongated vertically, overlapping, in a
single layer, 20-40 x 30-50 p, with outer walls smooth, 1. 5-2.0 p
thick, with inner walls closely marked by tubercles and very
short, irregularly oriented ridges, 4-5 p thick; aeciospores glo¬
bose, ovoid or ellipsoid, colorless, 21-28 x 21-34 p, mostly sub-
globose and about 24 x 28 p, finely and closely verrucose, with
the tubercles somewhat deciduous, one side partly smooth;
wall of aeciospore colorless, thin, scarcely 2 p thick including
tubercles, and less than 1 p without.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite
extent, punctate, immersed to slightly pustular, 90-250 p
wide by 100-125 p deep, flask-shaped in vertical section, open¬
ing outwards through a stoma of the overlying epidermis, with
the semblance of a short beak; peridium distinct; upper
peridial cells hyaline, isodiametric to irregular in transverse
section, 6-10 p across, of much greater depth, much over¬
lapped; lateral peridial cells radially elongated; walls of
peridial cells hyaline, thin; mouth of peridium surrounded by
slightly modified cells; uredospores colorless, white in mass,
few, extruded in delicate tendrils, short-stalked, irregularly
obovoid or ellipsoid, 15-27 x 27-51 p, averaging about 19 x
36 p; walls of spores thin, less than 1 p thick, with scattered,
short spines or spiny warts; three or four indistinct germ pores
more or less irregularly distributed, but often in pairs towards
the ends of the spores.
1932]
FAULL, THE GENUS MILESIA
71
III. Telia amphigenous, mostly hypophyllous, in over¬
wintered fronds, on indefinite, extensive brown areas, at times
involving entire fronds; teliospores within the epidermal cells,
very exceptionally within the guard cells, hyaline, rounded, or
irregular in outline and conforming to the shape of the con¬
taining epidermal cell, at times completely filling it, one- to
many-celled, with vertical septa, a single pore in the outer wall
of each cell; the cells of the teliospores with thin, smooth, col¬
orless walls, irregularly polygonal except along their free mar¬
gins, 8-16 x 12-25 h; basidiospores globular or subglobular,
7-8 n in diameter.
Hosts and Distribution:
O and I. Abies balsamea (L.) Mill., in Ontario.
II. Dryopteris marginalis (L.) A. Gray, in Ontario, Que¬
bec, New York, Massachusetts, New Hampshire.
III. Dryopteris marginalis (L.) A. Gray, in Ontario, New
York.
Type Locality: Lake Timagami, Ontario, on Abies balsamea and
Dryopteris marginalis. O, I, II and III.
Illustrations: Plate III, Figs. 12 a-d. — E. H. Moss in Ann.
Bot. 40: Text. Figs. 10, 11, 12, 21 F; Figs. 10, 31, 32 (1926). —
L. M. Hunter in Bot. Gaz. 83: Fig. 10 (1927).
Specimens Examined. —
O and I. On Abies balsamea.
Ontario: L. Timagami, various dates from July 29 — Aug. 22,
1924, /. H. Fault , Herb. J. H. Faull 4940, 4942, 4951, 4952,
7196, 7197, 7261, 7280, 7281, 7282.— Cultures, Aug. 1924,
J. H. Fault and W. R. Watson, Herb. J. H. Faull 7283-7299.
II. On overwintered fronds of Dryopteris marginalis.
Ontario: L. Timagami, May 18, 1924, May 12, 1925, June 14,
1926, J. H. Faull, Herb. J. H. Faull 4794, 7355, 7526 re¬
spectively.
Quebec: Hull, May 29, 1925, May 27, 1925, W. R. Watson, Herb.
J. H. Faull 7362, 7426 respectively. — Hudson, June 1913,
W. P. Fraser, Arthur Herb. 4481.
New Hampshire: Troy, Aug. 11, 1932, /. H. Faull.
Massachusetts: Becket, June 16, 1932, J. H. Faull and K. S.
Chester. — Mt. Greylock, June 17, 1932, /. H. Faull and K. S.
Chester.
New York: Labrador L., May 30, 1925, H. H. Whetzel, Myc.
Herb. N. Y. State Coll. Agr. 14732.
II and III. On overwintered fronds of Dryopteris mar¬
ginalis.
Ontario: L. Timagami, June 16, 1924, June 17, 1924, June 19,
1924, June 20, 1924 {type), June 25, 1924, June 30, 1924,
72
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
June 16, 1925, J. H. Fault , Herb. J. H. Faull 4862, 4865,
4869, 4877, 4896, 4904, 7370 respectively. — Cultures, Inocu¬
lations Aug. 22-27, 1927, collections June 29, 1928, on Dry-
opteris marginalis, E. H. Bensley, Herb. J. H. Faull 9845 a,
c, d, e, f, g. — Cultures, Inoculations Aug. 1924, collections
June 1925, on Dryopteris marginalis , W. R. Watson, Herb.
J. H. Faull 8261, 8263, 8269, 8276.
New York: Labrador L., May 30, 1925, H. H. Whetzel , Arthur
Herb. 4477. — Enfield Gorge, Ithaca, May 29, 1918, H. Fitz¬
patrick, Myc. Herb. N. Y. State Coll. Agr. 10755.
Milesia marginalis often occurs in great abundance through¬
out its range, to such an extent that in places it is difficult to
locate uninfected fern hosts. Though M. intermedia may be
prevalent in the same localities the two rusts are readily sep¬
arable, even on Abies. They differ with respect to their sper-
mogonia, frequency of uredinia, structure of uredinia, uredo-
spores, and incubation period on Abies.
The spermogonia of M. marginalis are larger in every way
than those of M. intermedia (27) and there are not nearly so
many of them on an individual needle. Unlike the latter, too,
a few are often to be found on the upper needle surface.
The uredinia are produced in great abundance, but they are
very minute. As in M. intermedia they are embedded in the
mesophyll and the overlying epidermis is not at all or but
slightly distended in pustular form. Hence they are more
easily located when found at the time of spore discharge.
Since the spores are comparatively large and the uredinia very
small it is not surprising to find that mature spores occur in
small numbers within a given uredinium at any one time. As
in other species of Milesia studied, so, too, in M. marginalis
the middle point of each uredinium occurs directly beneath a
stoma in the epidermis. The peridial wall consists of delicate
cells of relatively great depth, and they are much overlapped,
oriented towards the center of the dome. Those at the sides
are much more elongated still, and they represent mother col¬
umns, two or more along any radial line, from which spores
and peridial cells proper have not been developed. In the
young uredinia these columns are destined to become actively
sporogenous and each will contribute an elongated peridial cell
above. New columns peripheral to them develop in sequence.
Eventually the last columns formed are arrested in their de-
1932]
FAULL, THE GENUS MILESIA
73
velopment and never bear spores. At maturity the peridial
cells below the stoma elongate through the stoma, pushing the
guard cells laterally outward, so that a slightly differentiated
ostiolum in the form of a very short beak results. The mature
peridium showing this feature is correctly illustrated by Moss
(38, Text fig. 12).
The teliospores are produced in enormous quantities on
overwintered fronds. They are not formed, however, until
about the time the needles of Abies are unfolding. They then
germinate without a rest period, each basidium ordinarily bear¬
ing four delicate, spherical sporidia, 7 to 8 m in diameter. The
number of cells in a teliospore varies within wide limits; 4 to 8
are common, but as many as 25 have been counted.
The first reference to this species was made by Arthur (2,
5) under the name Milesia Kriegeriana who reported that
uredo material on Dryopteris marginalis had been sent to him
from Quebec, Canada, by Professor W. P. Fraser. I have had
the privilege of examining Fraser’s collection and find that it is
Milesia marginalis .
VI. SPECIES ON HISTIOPTERIS
19. Milesia Histiopteridis (G. H. Cunningham) Faull, n.
comb. II.
Milesina Histiopteridis G. H. Cunningham in Trans. N. Z.
Inst. 55:31 (1924).
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, aggregate or
scattered on discolored intercostal spots or on areas of in¬
definite extent, pustular, round, 0.15-0.4 mm. in width, cov¬
ered by brownish epidermis; peridium hemispheric, delicate,
closely adherent to the epidermis, opening at maturity along
with the overlying epidermis by an irregular apical pore;
peridial cells hyaline, flattened, isodiametrically to irregularly
polygonal, 7-15 m across; walls of peridial cells hyaline, thin,
less than 1 p thick; uredospores white, short-stalked, obovoid
or ellipsoid, 14-18 x 18-28 p, on an average about 16 x 24 p;
walls of spores colorless, thin, 0.7 -1.0 p thick, moderately and
finely verrucose.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
74
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
II. Histiopteris incisa (Thunb.) J. Sm. (P ter is incisa
Thunb.), in New Zealand.
III. Unknown.
Type Locality: Kelburn, Wellington, New Zealand, on Histiop¬
teris incisa. II.
Illustrations: Plate VIII, Figs. 31 a-d. — G. H. Cunningham in
Trans. N. Z. Inst. 55: Text. Fig. 101, and Plate I, Fig. 5 (1924).
Specimens Examined. —
New Zealand: Kelburn, Wellington, Aug. 14, 1922, E. H. Atkin¬
son , on Histiopteris incisa, Herb. G. H. Cunningham 1318
(II) ; part of type.
I am indebted to Professor J. J. Davis for access to the
mycological herbarium of the University of Wisconsin in
which I found material of Milesia Histiopteridis. Subsequent¬
ly a portion of the type was received from Dr. Cunningham.
In the original description of the rust Cunningham (9) cited
two collections, from Karori and Kelburn respectively, Wel¬
lington, N. Z., and he stated that “the rust is exceedingly com¬
mon in the localities where it has been collected, in fact,
scarcely a frond could be obtained free from the dead areas in
which the uredinia are embedded.” His description is accurate
except as to the peridium; this he misinterpreted. He thought
it consisted of “obovate, hyaline cells,” with their “outer walls
coarsely and densely verrucose.” As seen in vertical sections
the peridium is found to be typical for the genus. It is made
up of thin, delicate, flattened cells closely adherent to the over-
lying epidermis, following the contour of the latter, and in
periclinal sections its cells are coherent and polygonal. All of
the walls of the peridial cells are smooth and without markings.
VII. SPECIES ON NEPHROLEPIS
20. Milesia tenuis Faull, sp. nov. II.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregati, pustulati, rotundati,
0. 1-0.2 mm. diam., peridio ex cellulis hyalinis irregulariter
polygonalibus composite cincti; cellulae peridii 8-20 m diam.,
pariete 1.0- 1.7 m crasso. Uredosporae obovatae, ellipsoideae
vel subglobosae, subtiliter et densiuscule echinulatae, hyalinae,
12-17 x 20-28 |j, plus minusve circa 14 x 24 [ \ , episporio circa
1 m crasso. Teleutosporae ignotae.
1932]
FAULL, THE GENUS MILESIA
75
Hab. in foliis N ephrolepidis cordifoliae in Philippine Islands.
0 and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, aggregate or
scattered on brownish areas of indefinite extent, pustular,
round, 0.1 -0.2 mm. in width, covered by a dull brownish epi¬
dermis with centrally placed stomatic pore; peridium hemi¬
spheric, firm; peridial cells hyaline, isodiametrically to irregu¬
larly polygonal, 8-20 m across; walls of peridial cells hyaline,
1.0- 1.7 m thick; uredospores white, short-stalked, obovoid,
ellipsoid or subglobose, 12-17 x 20-28 m, on an average about
14 x 24 (j ; walls of spores colorless, thin, about 1 m thick, deli¬
cately and closely echinulate.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. N epkrolepis cordijolia (L.) Presl, in Philippine Islands.
III. Unknown.
Type Locality: Baguio, Luzon, Philippine Islands, on Nephro-
lepis cordijolia. II.
Illustrations: Plate VIII, Figs. 32 a-d.
Specimens Examined. —
Philippine Islands: Baguio, Benguit Prov., Luzon, on Nephro-
lepis cordijolia, Dec. 16-23, 1925, Mrs. M. S. Clemens
(7256 a), from Philippine Fungi deposited by E. D. Mer¬
rill in Univ. of Calif. Herb. 292430 (II); type.
21. Milesia columbiensis (Diet.) Arthur, in Mycologia, 7:
175 (1915), in part. II.
Milesina columbiensis Dietel in E. Mayor in Mem. Soc. Sci.
Nat. Neuchatel, 5(2):539 (1914). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, aggregate or
scattered on brownish areas of indefinite extent, pustular,
round, 0. 1-0.3 mm. in width, covered by yellowish brown epi¬
dermis with centrally placed stomatic pore; peridium hemi¬
spheric, very delicate; peridial cells hyaline, isodiametrically
to irregularly polygonal, 7-15 m across; walls of peridial cells
hyaline, 0.5 to 1.0 p thick; uredospores very short-stalked,
obovoid to ellipsoid, 14-23 x 25-37 m, on an average about 18 x
30 m; walls of spores colorless, thin, less than 1 n thick, deli-
76
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
cately and rather sparsely echinulate; occasional, strongly
spinose, club-shaped paraphyses, 12-15 m thick by 31-40 n long.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. N ephrolepis pendula (Raddi) J. Sm., in Colombia.
III. Unknown.
Type Locality: Quebradas, between Angelopolis and Guaca, Co¬
lombia, on N ephrolepis pendula. II.
Illustrations: Plate VII, Figs. 26 a-d. — P. Dietel in Mem. Soc.
Sci. Nat. Neuchatel, 5(2) : 5 5 9 , Fig. 67 (1914).
Specimens Examined. —
Colombia: Quebradas, August 1910, E. Mayor , on N ephrolepis
pendula (II); type.
Through the generous courtesy of Dr. Eugene Mayor I have
been enabled to examine the scanty, original material of this
rust, which up to the present is known from the one collection
only. My observations correspond closely to those of DietePs.
Thus as to the measurements of uredospores, Dietel gives
15-21 x 25-37 n, while my measures of 40 spores range from
14-23 x 25-37 m* While there is a centrally placed stomatic
pore above the dome of the uredinium, rupture may take place
by means of an irregular, elongated crack to one side of it.
A rust on Nephrolepis rivularis from Porto Rico, included
by Arthur (2, 5) under M. columbiensis is described in
this paper as M. insularis, n. sp.
22. Milesia insularis Faull, sp. nov. II.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, aggregati, pustulati, rotundati, 0.1-0.25 mm.
diam., peridio ex cellulis hyalinis irregulariter polygonalibus
composite cincti; cellulae peridii 7-15 m diam., pariete circa
1 m crasso. Uredosporae obovatae, ellipsoideae vel subglobo-
sae, subtiliter echinulatae, hyalinae, 15-23 x 20-30 m, plus
minusve 18 x 24 n, episporio 1-1.5 jj crasso. Teleutosporae
ignotae.
Hab. in foliis N ephrolepidis rivularis in Porto Rico.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, aggregate on
rather sharply defined small brown spots, pustular, round, 0.1
1932]
FAULL, THE GENUS MILESIA
77
to 0.25 mm. in width, covered by a yellowish brown epidermis
with centrally placed stomatic pore; peridium hemispheric,
colorless, delicate; upper peridial cells isodiametrically to
irregularly polygonal above, 7-15 m across; lateral peridial
cells radially elongate; walls of peridial cells hyaline, about 1
M thick; uredospores very short-stalked, obovoid, ellipsoid or
subglobular, 15-23 x 20-30 pi, on an average about 18 x 24 (j;
spore wall colorless, 1.0 to 1.5 jj thick, finely echinulate, the
points small.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. N ephrolepis rivularis (Vahl) Mett., in Porto Rico.
III. Unknown.
Type Locality: Agucaltaria, Porto Rico, on N ephrolepis rivu¬
laris. II.
Illustrations: Plate VII, Figs. 27 a-d.
Specimens Examined. —
Porto Rico: Agucaltaria, Nov. 25, 1913, F. L. Stevens (4855), on
N ephrolepis rivularis, Herb. Arthur, Plant ae Americae Sep-
tentrionalis (II); type.
Professor Arthur (2, 5) has included this rust under the
species M. columbiensis, the type of which was found on
N ephrolepis pendula in Colombia. Although the resemblance
is close there are, however, differences that warrant a separate,
specific designation. Thus the uredospores of M. insularis are
smaller than those of M. columbiensis, their walls are thicker,
and their spines are shorter and more abundant; likewise the
peridium is firmer and the uredinium is free from paraphysis-
like structures.
VIII. SPECIES ON PELLAEA
23. Milesia nervisequa (von Thiimen) Faull, n. comb. II.
Caeoma nerviscquum von Thiimen in Flora, 60:412 (1877).
II.
Milesina nervisequa (von Thiimen) Sydow in Monog. Ured.
3:481 (1915). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, aggregate on
greenish to dark brown areas of indefinite extent, sometimes
78
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
involving entire pinnae, or in elongate, rufous strips between
contiguous lateral veins, pustular, round, often confluent, 0.2
to 0.4 mm. in diameter, covered by yellowish epidermis which
finally ruptures at a centrally placed stomatic pore; peridium
hemispheric, colorless, delicate; peridial cells isodiametrically
to irregularly polygonal, 6 to 12 p across, with walls about 1 p
thick; spores hyaline, with finely granular contents, very short-
stalked, obovoid, ellipsoid or rarely subspherical, 14-18 x
20-31 p, averaging about 16 by 24 p; walls of spores 1.0 to
1.5 p thick, with numerous, scattered, minute, short spines.
III. Telia unknown.
Hosts and Distribution:
O and I. Unknown.
II. Pellaea hastata (Thunb.) Prantl, in Cape of Good
Hope, Natal, Transvaal, Madagascar.
Polypodium lycopodioides L., in Natal.
III. Unknown.
Type Locality: Somerset East, Cape of Good Hope, on Pellaea
hastata. II.
Illustrations: Plate VII, Figs. 28 a-d.
Exsiccati: Thuem. Myc. Univ. 1141.
Specimens Examined. —
A. On Pellaea hastata.
Cape of Good Hope: Somerset East, July 1876, P. MacOwan,
Thuem. Myc. Univ. 1141 (II); type.
Natal: Inanda, June 1881, J. M. Wood (441), Kew Herb. (Herb.
Mycol. M. C. Cooke, 1885) (II). — Durban, Apr. 13, 1911,
I. B. Pole-Evans, Myc. Herb. Union Dept, of Agr. 1370,
Pretoria, U. S. Afr. (II).
Transvaal: Lemana, Aug. 14, 1911, E. M. Doidge, Myc. Herb.
Union Dept, of Agr. 1836, Pretoria, U. S. Afr. (II). — Gars-
fontein, Nov. 28, 1910, P. J. Pienaar, Myc. Herb. Union
Dept, of Agr. 999, Pretoria, U. S. Afr. (II).
Madagascar: Tananarive, received at Herb. Crypt. Mus. Paris
May 21, 1919. M. Waterlot, Herb. Crypt. Mus. Paris (II).
B. On Poly podium lycopodioides.
Natal: Amanzimtoti, July 10, 1911, E. M. Doidge, Myc. Herb.
Union Dept, of Agr. 1634, Pretoria, U. S. Afr. (II).
According to von Thumen’s original description (46) the
uredospores of Milesia nervisequa are hyaline. If by this
white was meant, then the uredinia considered apart from the
unknown telia are those of a Milesia. The uredospores as I
1932]
FAULL, THE GENUS MILESIA
79
have seen them in the dried specimens available are colorless.
They may possibly be white, but a final statement awaits
observation on fresh material. Until the spore color is finally
determined the species may very well be left in the genus
Milesia. The epidermis overlying the uredinia is yellowish,
but the color is wholly in the epidermal cells themselves. Von
Thiimen overlooked the peridium, which is well developed and
can be easily demonstrated especially in the younger uredinia.
He stated that the spores were catenulate and that they were
smooth, in both of which points he was in error.
The specific name given by von Thiimen is not as inappro¬
priate as Sydow (44) maintains. It is true that the lesions
often bear no apparent relationship to the veins, but it also
happens that they are sometimes restricted laterally between
adjacent veins, quite striking from their bright rufous color,
especially as manifested on the upper leaf surface; and the
uredinia are then distributed in somewhat lineal series between
and parallel to the veins. But even in such cases the lesions
may and frequently do extend beyond their original bounds,
involving areas of indefinite extent and without relation to the
veins of the affected leaf. The lesions, it may be noted in
passing, bear uredinia while still green or greenish and they
become brown quite tardily.
Dr. E. M. Doidge (14) lists several collections of M. nervi-
sequa from Natal and the Transvaal. She states that the ure-
dospores are hyaline, and that the uredinia long remain closed.
A record of still wider distribution is afforded by a Madagascar
collection received from the Museum National d’Histoire
Naturelle, Paris, through the courtesy of Dr. Roger Heim.
This material, in excellent condition, was provisionally deter¬
mined by N. Patouillard as “Hyalopsora Waterloti n. sp. (ad
int.) sous les feuilles d’un Pellaea (cfr. P. viridis ).” The rust
is identical with M. nervisequa (von Thiimen) Faull; and the
host appears to be Pellaea hastata, a name synonymous with
the preferable name P. viridis (Forskal) Prantl according to
Christensen’s Index Filicum (1906).
One of Dr. Doidge’s collections is of exceptional interest, a
collection of rusted Polypodium lycopodioides. I have exam¬
ined the rust on this fern with a great deal of enthusiasm be¬
cause it is the first known instance of a supposedly Milesia
80
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
rust on a host of the large Pleopeltid section of the genus Poly¬
podium. Uredinia only were found, and their morphological
characters both as to spores and peridia correspond closely to
those of M. nervisequa. My description of the rust in ques¬
tion is as follows. — “Uredinia hypophyllous, subepidermal,
scattered or loosely aggregate on dark brown or greenish areas
of indefinite extent, pustular, round, 0.1 to 0.4 mm. in diam¬
eter, covered by yellowish brown epidermis which finally rup¬
tures at a centrally placed stomatic pore; peridium hemi¬
spheric; peridial cells hyaline, isodiametrically to irregularly
polygonal, 6-11 x 9-15 m; walls of peridial cells hyaline, 1.0-
1.5 m thick; uredospores hyaline, short-stalked, obovoid, ellip¬
soid or subglobose, 13-19 x 19-33 p, averaging about 16 x 26 p;
walls of spores 1.0- 1.3 p thick, with minute, short, numerous,
scattered spines, rarely also with an occasional wart or longer
spine.” If this description be compared with the one given
above for the species, drawn entirely from a study of the rust
on Pellaea hastata, it will be seen that the two are essentially
identical. My conclusion is that until further collections have
been made, other stages examined, and such experimentation
as cross inoculations carried out, the wisest course is to diag¬
nose Dr. Doidge’s one collection on Polypodium lycopodioides
as Milesia nervisequa. Granted that such diagnosis be cor¬
rect, M. nervisequa then would constitute the first known case,
according to my interpretation, of a Milesia infecting ferns be¬
longing to two distinct genera.
Dr. Doidge reported the rust on Polypodium lycopodioides
(Bothalia, 2:162. 1927) under the name Milesina Dieteliana
(Syd.) Magnus. She appended a description of uredinia and
teliospores that appears to be largely drawn from Sydow (44)
and applicable mainly to Milesia Polypodii White [Milesina
Dieteliana (Syd.) Magnus] as known in Europe on Poly po¬
dium vulgare. The chief point of concern in this connection
is as to whether or not Dr. Doidge actually found teliospores
in the South African collection on P. lycopodioides. She kind¬
ly shared a portion of her collection with me for my study; but
in this I find uredinia only. I may add that this rust on P.
lycopodioides proves to be distinctly different from Milesia
Polypodii in that its uredospores are much smaller than those
of the latter and more finely spinulose, and its peridial cells
1932]
FAULL, THE GENUS MILESIA
81
have thicker walls. If now the teliospores can be found the
rust on P. lycopodioides affords opportunity for a fascinating
life history study, partly because of its distribution far beyond
the limits of Abies, and also because its host belongs to a sub¬
genus of Polypodium quite distinct from the subgenus which
includes the host of M. Polypodii.
Finally, the finding of species of Milesia, such as M. nervi-
sequa, M. Histiopteridis, etc., in regions as far removed as
South Africa, New Zealand, the Philippines, South America
and the West Indies, assuming that they are phylogenetically
related to the northern species, leads one to surmise that many
more remain yet to be discovered, especially in the southern
hemisphere. One may also be justified in speculating on the
likelihood of an evolution of the genus on the Filicales quite
independent of an alternate host.
IX. SPECIES ON POLYPODIUM
24. Milesia Polypodii B. White in Scot. Nat. 4:162 (1877).
(II). II and III.
Melampsorella Dieteliana Sydow in Annal. Mycol. 1:537
(1903). II and III.
Milesina Dieteliana (Syd.) Magnus in Ber. Deutsch. Bot.
Ges. 27:325 (1909). II and III.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brown areas of indefinite ex¬
tent, pustular, round, 0. 1-0.2 mm. in width, covered by a
brownish discolored epidermis with centrally placed stomatic
pore; peridium delicate but firm; peridial cells hyaline, iso-
diametrically to irregularly polygonal, 8-18 p across, lateral
ones more or less radially elongated; walls of peridial cells
hyaline, 1 p or less thick; uredospores colorless, short-stalked,
obovoid to ellipsoid, sometimes subglobose, 15-26 x 24-48 p,
on an average about 19 x 35 p; walls of spores 1-2 p thick,
diffusely and rather strongly echinulate.
III. Telia on overwintered fronds, hypophyllous, on in¬
definite, extensive brown areas; teliospores within the epi¬
dermal cells, occasionally in the guard cells, hyaline, rounded,
or irregular in outline and conforming to the shape of the con¬
taining epidermal cell, often completely filling it, one- to many-
82
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
celled (up to fifty-celled), with anticlinal septa, a single pore in
the outer wall of each cell; the cells of the teliospores with
thin, smooth, colorless walls, irregularly polygonal except
along their free margins, 8-20 x 12-23 p; basidiospores sub-
globular, 7 to 9 p in diameter.
Hosts and Distribution:
O and I. Unknown.
II. Polypodium vulgare L., in Great Britain, Germany,
Switzerland, Austria, Denmark, Norway, Poland, Spain, France,
Czechoslovakia, Caucasus.
Polypodium vulgare L. var. serratum , in Scotland, accord¬
ing to Grove (21).
III. Polypodium vulgare L., in Great Britain, Germany,
France, Switzerland, Poland, Czechoslovakia.
Type Locality: Reichip, Perthshire, Scotland, on Poly podium
vulgare. II and III. (See comments below on finding teliospores in
the type material.)
Illustrations: Plate I, Figs. 1 a-d. — White in Scot. Nat. 4:163,
Fig. 5 (1877). — Grove in Brit. Rust Fungi, Fig. 281, p. 376 (1913).
— Kamei in Trans. Sapporo Nat. His. Soc. 12(1) : 29, Fig. 2 B
(1931). — R. Gonzalez Fragoso in Flora Iberica, Uredales, II, Fig.
139, p. 283 (1925).
Exsiccati: Syd. Myc. Germ. 62, under name Melampsorella
Dieteliana Syd., 1328. — Syd. Ured. 1650, under name Uredo Scolo-
pendrii (Fuckel) Schroet., 2396, under name Melampsorella Die¬
teliana Syd., 2597. — Vestergren Microm. Rar. Sel. 1309, under
name Uredo Scolopendrii (Fuckel) Schroet., 1702, under name
Milesina Dieteliana (Syd.) Magn. (most of synonyms cited on label
are incorrect). — Krieg. Fg. Saxon. 817, under name Uredo Scolo¬
pendrii (Fuckel) Schroet.
Specimens Examined, all on Poly podium vulgare. —
Scotland: Perthshire, July 3, 1877, F. Buchanan White (II, III);
part of type. — Reichip, Perthshire, June 1877, F. Buchanan
White (II); part of type. — Glen Tilt, Perthshire, Aug. 1877,
F. Buchanan White (II); part of type.
England: Lytchett Minster, Dorset Co., Nov. 26, 1931, G. D.
Darker (II).
Germany: Schmilka, Saxony, Aug. 26, 1903, H. and P. Sydow, Syd.
Myc. Germ. 62 (II). — Schmilka, Saxony, Sept. 28, 1895,
G. Wagner, Syd. Ured. 1650 (II). — Polenzthal, Saxony, Aug.
26, 1891, W. Krieger, Krieg. Fg. Saxon. 817 (II).
Norway: Insula Anuglen, June 12, 1907, T. Vestergren, Vestergren
Microm. Rar. Sel. 1309 (II).
France: Potigny, April 14, 1911, R. Maire, Syd. Ured. 2396 (II). —
Wagenberg bei Wasselnheim, Alsace, Oct. 23, 1914, A. Lud¬
wig, Syd. Myc. Germ. 1328 (II). — Axat, Aude, Apr. 30, May
1 and 3, 1932, G. D. Darker (II, III).
1932]
FA ULLr THE GENUS MILESIA
83
Switzerland: Neuchatel, Oct. 20, 1913, E. Mayor, Syd. Ured. 2597
(II).— Neuchatel, Nov. 5, 1909, E. Mayor (II). — Tete
Plumee sur Neuchatel, Nov. 5, 1909, E. Mayor, Vestergren
Microm. Rar. Sel. 1702 (II). — Neuchatel, June 12, 1932,
G. D. Darker, overwintered fronds (II, III).
Poland: Ojcow, Krakow, 1913, A. Wroblewski (II). — Ojcow, June
27, 1924, Wanda Konopacka, Herb. W. Siemaszko (II, III).
— Ojcow, Aug. 26, 1925, W. Konopacka (II).
Czechoslovakia: Kralovany, Slovakia, July 27, 1932, G. D. Dar¬
ker, overwintered fronds (II, III).
Caucasus: Suchum, Oct. 25, 1914, W. Siemaszko (II).
Milesia Polypodii was first described by Dr. F. Buchanan
White (48). His type materials, portions of which I have had
the opportunity of examining through the courtesy of Mr. J.
Ramsbottom, were deposited in the Mycological Herbarium of
the British Museum of Natural History as part of a bequest
from C. E. Broome in 1886. The finding of this colorless rust
greatly interested Dr. White, and before describing it speci¬
mens both fresh and dried were submitted to Miles Joseph
Berkeley and C. E. Broome. Both of these men made careful
examination of the specimens and there was quite an extensive
correspondence between them and White on the subject. The
correspondence, copies of which I have seen through the cour¬
tesy of the Director of the British Museum, contained de¬
scriptions of the fungus, drawings of the peridium and uredo-
spores, and expressions of opinion as to the nature of the
fungus. Berkeley was inclined to regard it as one of the Fungi
Imperfecti, nearer to Sphaeropsis than anything else; to White
it was a rust recalling to his mind the Uredo on Scolopendrium
(now known as Milesia Scolopendrii) ; to Broome it was of
curious interest, leading him to surmise that it might come
close to Endophyllum. With all of this information available
it is exasperating that P. Magnus (37) should remark that we
do not know on what fungus White based his genus Milesia and
his description of M. Polypodii, comments to be re-echoed later
by Grove (21).
After going over the type material I find that White’s de¬
scription was accurate, discriminating and carefully illustrated.
It speaks highly for his care in observation that he detected a
peridial covering of the uredinium, that he saw the minute
apical ostiole in the papilliform uredinium, and that he was
84
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
impressed by the white spores. That the uredospores of this
species are white, and that there might be any generic sig¬
nificance in that phenomenon, remained unemphasized and
usually unnoted for considerably more than a quarter of a cen¬
tury after. It speaks highly for White’s acumen that he in¬
terpreted this fungus as a rust, at a time when most
mycologists would have referred it to the imperfect fungi. His
description is quoted as follows: —
“Milesia B. White, g. n.”
“Peridium endophyllum reticulatum, basi inter cellulas matricis
radicante; sporae obovatae echinulatae ex ostiolo minuto demum
ejectae.
“Allied to Endophyllum Lev. Peridium minute, reticulated, im¬
mersed in the substance of the matrix, into which the base sends
root-like fibres. Spores distinctly spiny, ejected copiously from a
minute ostiolum at the summit of the peridium.
“It is with very great pleasure that I dedicate this genus to the
Rev. Miles Joseph Berkeley, M.A., F.L.S., to whose researches our
knowledge of the mycology, not only of Britain, but of every part
of the world, is so deeply indebted, and whose kindness to all stu¬
dents of the same branch of botany is as well known as it is un¬
failing.
“Milesia polypodii B. White sp. n.”
“M. hypogeneum; peridiis sphaericis minutis, pallidis, dispersis;
sporis albidis obovatis echinulatis.
“On the underside of living but fading leaves of Poly podium vul¬
gar e in June and July. First found in the den of Reichip; then in
other parts of Perthshire.
“Spores .0012-.0017 inch long” (i. e. 30-42 |j long). “(Plate II.
Fig. 5). The peridia are scattered in an irregular manner on the
underside of the frond of the fern, and look under the lens like little
pallid mammiform projections very slightly elevated. The spores
are ejected copiously from the ostiolum, and from their whitish color
and abundance are very perceptible. The spores themselves appear
under the microscope to be filled with a granular matter.”
It will be noted that the only spores described by White
were the uredospores. That the teliospores might be found in
the unmodified epidermal cells was not surmised by anyone at
that time. An examination of White’s material, however, re¬
veals the fact that they were present in some of it, and that
there were basidia and basidiospores as well. My description
of the teliospores as given above, is drawn from the type,
specimens of overwintered fronds collected July 3, 1877.
1932]
FAULL, THE GENUS MILESIA
85
Sydow (42) discovered M. Polypodii many years later in
Germany (1903), and apparently unaware that it had been
previously studied and named, described it as a new species
under the name Melampsorella Dieteliana. Sydow found and
recognized teliospores in his specimens, and noted that the
uredospores were hyaline. It was six years later (1909) that
Magnus (37), discounting and discarding White’s work, ac¬
cepted Sydow’s less discriminating description, and transferred
the species to his inaccurately conceived genus Milesina.
As nineteen collections, some of them from widely separated
regions, have been available to me in my study of M. Poly¬
podii I have taken the opportunity of making a careful com¬
parison of their uredospores, partly to determine what may be
regarded as approximately correct average measurements, and
partly to secure data illustrative of the range of variability
manifested by the uredospores of a single species of Milesia.
The appended table is a representative presentation of the
results obtained. Forty spores, taken at random, were meas¬
ured from each of the seven collections listed below.
TABLE 6
MEASUREMENTS OF UREDOSPORES OF M. POLYPODII FROM VARIOUS
COLLECTIONS
Source of
Material
Scotland (type)
Switzerland
Switzerland
Poland
France
Germany
Norway
Average
Breadth x
Length
17-20 x 23-40 fx
17- 23 x 26-40
18- 26 x 30-48
17-23 x 28-48
16- 20 x 26-48
17- 26 x 26-45
16-23 x 27-45
16-26 x 23-48 p.
Average Breadth
x Length
18.7 x 31.0 p,
17.9 x 38.9
18.5 x 35.4
19.2 x 32.2
20, .0 x 36.7
20.4 x 31.9
22.1 x 37.0
19.5 x 34.7 p,
Ratio of
Breadth to Length
1:1.66
1:2.17
1:1.91
1:1.68
1:1.84
1:1.56
1:1.68
1:1.78
Everyone who has looked at the uredospores of a Milesia is
familiar with the fact that the mature spores of any one ure-
dinium manifest enormous variations of shape, size and rela¬
tive proportions. It is further clear, especially as revealed by
the foregoing table, that collections from various localities may
exhibit rather surprising differences in average sizes and pro¬
portions. What may be the explanation of these latter phe¬
nomena is not clear, as for example, whether or not there may
be morphologically distinct strains of such a species as M.
86
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Polypodii just as there are strains of Taraxacum officinale or
Draba verna in the seed plants. Before a reasoned answer
could be given it would be necessary to observe the spore out¬
put from the same mycelia throughout their lifetime, to make
comparisons of the spores of the same species of rust from
different localities under identical climatic conditions, and to
determine whether or not the host exercises an influence — all
of which could quite readily be ascertained. Obviously it is
important in making spore measurements to make sure that
the spores are fully mature.
It would not be amiss at this point to recall a confusion asso¬
ciated with Milesia Polypodii that long prevailed in the litera¬
ture, and which was the result of a careless transcript from
White’s original paper. I refer to repeated references to a
supposedly existent, but really non-existent, Milesia Polygoni
on Polygonum viviparum (see Annals and Magazine of Natu¬
ral History, 5th Series, 1:27. 1878 — the source of the mistake;
Hedwigia, 17:83. 1878; Saccardo’s Sylloge Fungorum, 7(2):
768. 1888; Ber. Deutsch. Bot. Ges. 27:325. 1909). This
error was pointed out and corrected in Grevillea, 17:16 (1889).
In pointing out the error another was made in ascribing the
authorship of M. Polypodii to Berkeley and White.
The complete life history of M. Polypodii remains yet to be
elucidated, though it would appear that such could be done
quite easily. The rust is widely distributed in Europe, not
rare, and its teliospores occur abundantly on overwintered
fronds. Almost certainly it will be found that Abies is the
alternate host and it is to be anticipated that the aecial stage
runs a rapid course on needles of the current season. As noted
elsewhere in this paper two other species of Milesia on
species of Poly podium have been demonstrated to have Abies
as the alternate host.
By way of conclusion I venture to express the opinion that
when all the facts are known the evolution and the distribution
of Milesia on Polypodium will prove to be a topic of intense
and significant interest. Polypodium is apparently a favorable
host for Milesia, and search for this rust on the many species
of this large genus promises a rich yield. A good beginning
has been made. We know that M . Polypodii is frequent on P.
vulgare throughout a wide range in Europe and southward to
1932]
FAULL, THE GENUS MILESIA
87
the Caucasus in Asia (W. Siemaszko in Trans. Suchum Gar¬
den Field Sta. 2:9. 1915). Kamei (30) has just described M.
jezoensis on the same host in Japan. M. laeviuscula occurs in
two forms, typically so distinct as to be fairly recognizable as
distinct species, on P. calif ornicum and P. glycyrrhiza respec¬
tively, in the western part of North America. M. polypodo-
phila, a remarkable representative of the species, occurs on
P. virginianum in the eastern part of North America.
25. Milesia jezoensis (Kamei & Hiratsuka) Faull, n. comb.
O, II and III.
Milesina jezoensis Kamei & Hiratsuka in S. Kamei in Trans.
Sapporo Nat. His. Soc. 12, 1:32 (1931). O, II and III.
O. “Spermogonia hypophyllous, scattered on yellowish dis¬
colored areas, subcuticular, more or less globose, 110-165 p in
breadth, 110-130 p in height; spermatiophores obclavate, sep¬
tate; spermatia oblong, 6. 5-9.0 p long, 2.0-3.0 p broad.”
(From cultures. Kamei, 30).
I. Not yet recognized, but certainly existent.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely aggregate on greenish to brown areas of indefinite ex¬
tent, pustular, round, 0.2 to 0.4 mm. in diameter, covered by
yellowish brown epidermis which finally ruptures at a centrally
placed stomatic pore; peridium hemispheric, firm, readily sep¬
arable from overlying epidermis; peridial cells colorless, iso-
diametrically to irregularly polygonal, 7-11 x 8-15 p, lateral
cells often radially elongate; walls of peridial cells 1.0 to 1.2 p
thick; uredospores white, very short-stalked, obovoid, ellip¬
soid to subspherical, sometimes irregular in shape, 16-22 x
25-47 p, averaging about 19 x 35 p; spore walls very finely
verruculose, especially at the apex, 0.7 to 0.9 p thick.
III. Telia on overwintered fronds, mostly hypophyllous,
on brown areas of indefinite extent; teliospores within the epi¬
dermal cells, hyaline, rounded, or irregular in outline and more
or less conforming to the shape of the containing epidermal
cell, often completely filling it, one- to many-celled, up to
twelve or more, with vertical septa, a single pore in the outer
wall of each cell; the cells of the teliospores with thin, smooth,
colorless walls, irregularly polygonal except along their free
margins, 8-19 x 8-27 p.
88
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Hosts and Distribution:
O. Abies Mayriana Miyabe and Kudo, in Japan. (Cul¬
tures).
II and III. Polypodium vulgare L., in Japan.
Type Locality: Lake Shikaribetsu, Prov. Tokachi, Japan, on
Poly podium vulgare. II and III.
Illustrations: Plate VIII, Figs. 33 a-d. — Kamei in Trans. Sap¬
poro Nat. His. Soc. 12, 1:27-33, Figs. 1, 2 C, 3 (1931).
Specimens Examined, all on Polypodium vulgare. —
Japan: Lake Shikaribetsu, Prov. Tokachi, Sept. 20, 1930, S. Kamei
(II). — Lake Shikaribetsu, Prov. Tokachi, June 11, 1931,
S. Kamei, overwintered fronds (II, III); type locality.
An opportunity to examine both uredinial and telial ma¬
terials of Milesia jezoensis has been afforded through the
courtesy of Dr. S. Kamei. It is from these materials that my
descriptions of uredinia and telia have been drawn.
Among the species of Milesia now known to occur on
ferns of the genus Poly podium, M. jezoensis occupies an inter¬
mediate position between M. Polypodii {Milesina Dieteliana)
and M. laeviuscula forma glycyrrhiza , so far as can be judged
from the morphological characters of the uredospores and the
uredinial peridia. The uredospores of the former are strongly
spinulose, those of the latter are smooth, while those of M.
jezoensis are densely verruculose at their upper ends but al¬
most smooth elsewhere. The characters of the uredinial
peridium of M. jezoensis likewise are intermediate, both as to
firmness of the peridium and the thickness of its cell walls.
An allocation of these species places M. Polypodii at one
end of the series and M. polypodophila distinctly apart at the
other end. Incidentally it may be surmised that the spermo-
gonia are also likely to exhibit a gradation, as indicated by the
differences manifested in M. jezoensis and M. polypodophila,
the only two of them whose spermogonia are known.
The foregoing serial arrangement of these species of
Milesia on Polypodium suggests an evolutionary sequence. As
such it emphasizes the pronounced mutability of the uredinium
and of the spermogonia, a feature so generally evident through¬
out the genus Milesia. In this connection the geographical
distribution of the species under consideration is of particular
interest. Following the series through we note that M. Poly¬
podii occurs in Europe and ranges eastward towards and even
1932]
FAULL, THE GENUS MILESIA
89
into continental Asia; M. jezoensis is found in Japan in east¬
ern Asia; M. laeviuscula forma glycyrrhiza on P. glycyrrhiza
ranges from Alaska in western America southward as far as
Oregon; M. laeviuscula forma typica, so close to its sister
form, occurs in the contiguous state of California on P. cali-
fornicum, a host scarcely distinguishable from P. glycyrrhiza;
and M. polypodophila is known only from eastern North
America. For comments on M. nervisequa on P. lycopodioides,
see p. 79.
We owe our knowledge of the life history of M. jezoensis to
Kamei (30). Kamei found telial material for his experimen¬
tation in early June on overwintered fronds. The teliospores
soon germinated, producing typical basidia — “cylindrical to
clavate, four-celled, about 45 m long and 10 m wide; sporidia
subglobose, about 11 m long x 7 m wide.” Kamei made sowings
of basidiospores on potted plants in a greenhouse. Infection
took place on the new needles, spermogonia appearing about
twenty days after inoculation. Unfortunately the infected
needles dropped off prematurely and without any development
of aecia — an experience that can probably be avoided under
artificial conditions by more careful attention to the vigor of
the experimental host plants and their exacting requirements.
Kamei ’s experiment, though convincing, was incomplete; it
should be repeated and the aeciospores when obtained inocu¬
lated on to the fern host.
26. Milesia polypodophila (Bell) Faull, n. comb. O, I, II
and III.
Uredinopsis polypodophila Bell in Bot. Gaz. 77:25 (1924).
II.
Peridermimn pycnogrande Bell in Bot. Gaz. 77:24 (1924).
O and I.
Milesia pycnograndis (Bell) Arthur in North Am. Flora,
7:685 (1925). O, I, II and III.
Milesina polypodophila (Bell) Faull in Proc. Int. Congr. PI.
Sci. (Ithaca, 1926) 2:1736 (1929). O, I, II and III.
O. Spermogonia on needles 3 to 9 years old, hypophyllous,
few, in two rows, one on each side of midrib, forming distinct
but not conspicuous, light-colored areas, subepidermal, deep-
seated, globoid, 175 to 250 m broad by 175 to 243 m deep,
averaging about 200 m broad by 212 m deep; apical pore a
stoma or a short slit, 7-12 x 19-29 n, parallel to the long axis of
the needle; spermatiophores unbranched, septate, producing
90
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
the spermatia catenulately; spermatia hyaline, unicellular,
narrowly ellipsoid, 1. 7-3.0 x 5.0-7.0 p.
I. Aecia hypophyllous, on needles 3 to 9 years old, in two
rows, white, cylindrical, 0. 2-0.3 mm. in diameter by 0.4 to 1.0
mm. high; peridium colorless, thin, firm, rupturing at the
apex; peridial cells polygonal, elongated vertically, much over¬
lapping, in a single layer, 15-27 x 22-38 p exclusive of laps,
up to 55 p in total length, with outer walls smooth, 1.0- 1.5 p
thick, with inner walls closely and finely marked by delicate
warts and short ridges, 3-4 p thick; aeciospores white, subglo-
bose or short ellipsoid, 18-27 x 21-30 p, averaging about 25 x
26 p, finely and closely verrucose; walls of aeciospore thin,
1.0- 1.5 p thick.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite ex¬
tent, pustular, covered by pale greenish or brownish epidermis,
0.2 to 0.5 mm. wide by 0.10 to 0.16 mm. deep, rupturing at a
centrally placed stoma in the overlying epidermis; peridium
distinct, firm, mostly separate from the epidermis; peridial
cells hyaline, polygonal, isodiametric or irregular, 8-12 x
8-20 p, walls about 1.5 p thick; uredospores colorless, white in
mass, abundant, extruded in delicate tendrils, short-stalked,
fusiform-obovoid or fusiform, acute or acuminate at apex,
occasionally forked at apex, narrowed below, 13-21 x 38-68 p,
averaging about 17-20 x 51-53 p; walls of spores thin, about
1 p thick, smooth, with 4 to 6 germ pores in pairs towards poles
and at equator.
III. Telia on overwintered fronds, hypophyllous, on in¬
definite, extensive brownish areas; teliospores within the epi¬
dermal cells, common in the guard cells, hyaline, rounded, or
irregular in outline and conforming to the shape of the con¬
taining epidermal cell, usually completely filling it, 3- to 50-
celled, commonly 20- to 30-celled except in the guard cells
where the number is 4 to 7, with anticlimal septa, a single pore
in the outer wall of each cell; the individual cells of the telio¬
spores with thin, smooth, colorless walls, irregularly polygonal
except along their free margins, 7-15 x 8-22 p.
Hosts and Distribution:
O and I. Abies balsamea (L.) Mill., in Ontario, Quebec,
Nova Scotia, Maine, Vermont, New Hampshire.
1932]
FAULL, THE GENUS MILESIA
91
II. Polypodium virginianum L., in Ontario, Quebec, Nova
Scotia, Maine, New Hampshire, Vermont, Massachusetts,
Connecticut, New York, Pennsylvania, Tennessee.
III. Polypodium virginianum L., in Ontario, Quebec, Nova
Scotia, New Hampshire, Vermont, Massachusetts, Connecticut,
New York, Pennsylvania.
Type Locality: Lake Timagami, Ontario, on Abies balsamea and
Poly podium virginianum. O, I, II and III.
Illustrations: Plate I, Figs. 4 a-d; Plate IX, Figs. 36 and 37. —
E. H. Moss in Ann. Bot. 40: Text Figs. 8, 9, 21 E; Figs. 9, 33, 34,
35 (1926).— L. M. Hunter in Bot. Gaz. 83: Fig. 11 (1927).— S.
Kamei in Trans. Sapporo Nat. His. Soc. 12, 1:29 Fig. 2 A (1931).
Specimens Examined. —
O and I. On Abies balsamea.
Ontario: L. Timagami, various dates, July 12 to Sept. 1, 1921 to
1928, J. H. Faull and H. P. Bell, Herb. J. H. Faull 4329,
4843, 5952 {type), 6022, 7194, 8050, 8380, 8596, 8597,
8598. — Cultures, inoculations July 1924, collected Aug. 1927
and Aug. 1928, J . H. Faull and W. R. Watson, Herb. J. H.
Faull 8596-8598.
Quebec: Kenogami, July 25, 1923, Aug. 26, 1923, J. FL. Faull, Herb.
J. H. Faull 5477, 4632 respectively.
Nova Scotia: Liverpool, July 23, 1929, J. H. Faull, Herb. J. H.
Faull 9265. — Creeper’s Lake, July 31, 1930, J. H. Faull.
Maine: Oquossoc, May 17, 1931 (from season of 1930), J. H.
Faull. — Oquossoc, Oct. 25, 1931 (overmature), J. FL. Faull.
Vermont: Searsburg, June 17, 1932 (immature), J. H. Faull and
K. S. Chester. — Sherburne Pass, Aug. 10, 1932, J. H. Faull.
New Hampshire: Mt. Jefferson, Aug. 31, 1932, /. H. Faull.
II. On Polypodium virginianum.
Ontario: Lake Timagami, July 27, 1924, Aug. 1, 1924, J. FL. Faull,
Herb. J. H. Faull 4938, 7195, 7334.
Maine: Oquossoc, May 17, 1931, J. H. Faull, Herb. J. H. Faull
9265, overwintered fronds but teliospores not yet formed. —
Oquossoc, Oct. 25, 1931, /. H. Faull, Herb. J. H. Faull
10402.
Vermont: Warren, July 15, 1932, J. H. Faull and K. S. Chester. —
Sherburne Pass, Aug. 10, 1932, J. H. Faull.
New York: Whitehall, Aug. 10, 1932, /. H. Faull. — Warrensburg,
Aug. 8, 1932, /. H. Faull.
Tennessee: Gatlinburg, June 20, 1931, FL. FL. Whetzel et al., Myc.
Herb. N. Y. State Coll, of Agr. 19335, overwintered fronds.
II and III. On Polypodium virginianum.
Ontario: Lake Timagami, Aug. 22, 1922, June 19, 1924, June 19,
1924, June 22, 1924, June 27, 1924, J. H. Faull and H. P.
Bell, Herb. J. H. Faull 6788, 4867, 4870 {type), 4875, 4899,
overwintered fronds. — Cultures, inoculations Aug. 9, 1924,
collected July 5, 1925, and July 21, 1925, J. FL. Faull and
92
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
W. R. Watson, Herb. J. H. Faull 7374 (Expmts. 212-215),
7375 (Expmts. 216-218) (II, III). — Inoculations Aug.
1924, collected June 1925, /. H. Faull and W. R. Watson,
Herb. J. H. Faull 8262, 8264, 8266 (II).
Quebec: Kenogami, Aug. 26, 1923, /. H. Faull, Herb. J. H. Faull
4633, overwintered fronds.
Nova Scotia: Liverpool, July 23, 1929, /. H. Faull, Herb. J. H.
Faull 9265, overwintered fronds. — Creeper’s Lake, July 31,
1930, J. H. Faull, overwintered fronds.
New Hampshire: Troy, Aug. 11, 1932, /. H. Faull.
Vermont: Searsburg, June 17, 1932, J. H. Faull and K. S. Chester.
Massachusetts: Becket, June 16, 1932, J. H. Faull and K. S.
Chester. — Guilder Pond, Mt. Washington Tp., July 13, 1932,
J. H. Faull and K. S. Chester.
Connecticut: Sage’s Ravine, Salisbury, June 17, 1926, G. P. Clin¬
ton and W. R. Hunt, Herb. Conn. Agr. Expmt. Sta. — North
Canaan, June 17, 1926, G. P. Clinton and W. R. Hunt, Herb.
Conn. Agr. Expmt. Sta. — People’s Forest, Barkhamstead,
June 17, 1926, G. P. Clinton and W. R. Hunt, Herb. Conn.
Agr. Expmt. Sta.
New York: Lake Placid, Aug. 10, 1932, J . H. Faull.
Pennsylvania: Howard, May 30, 1927, H. W. Thurston, Jr., Herb.
J. H. Faull 8429, overwintered fronds.
The peridermia of Milesia polypodophila were first found by
J. H. Faull and H. P. Bell on Abies balsamea, Aug. 3, 1921
(Herb. J. H. Faull 5952). The uredinia were located by Bell
in the same region (L. Timagami, Ontario) during the sum¬
mer of 1922. Bell (6) described these stages respectively
under the names Peridermium pycno grande and Uredinopsis
polypodophila. The telia were discovered by Faull in the
Timagami region on overwintered fronds of Poly podium vir-
ginianum, June 19, 1924 (Herb. J. H. Faull 4867). With the
assistance of W. R. Watson culture experiments were at once
undertaken, with the result that Bell’s rusts on the Fir and on
the Polypody were demonstrated to be phases of the one spe¬
cies. Cunningham (The Rust Fungi of New Zealand, 1931,
p. 54) makes the unwarranted and impossible statement that
Bell (1924) first established “the connection between an aeci-
diosorus on Abies balsamea and uredosori of Milesina poly¬
podophila on Poly podium vulgare.” Bell attempted no experi¬
mentation whatever in this connection.
Milesia polypodophila has proved to be of exceptional in¬
terest both morphologically and biologically, for it stands
1932]
FAULL, THE GENUS MILES I A
93
apart in many of its characteristics from all other known spe¬
cies of the genus.
Milesia polypodophila is perennial in the aecial host and an
infection once established is apparently capable of continuing
for an indefinite period of years. It exercises a mild, stimu¬
lating effect on its host, in that some of the dormant buds are
forced; the result is a loose type of broom. There is appar¬
ently, however, no interference with the normal, physiological
habits of the foliage of the current season, and there is never
a devastation of great numbers of the older needles; so these
brooms live on from year to year, slowly becoming larger, but
not wholly exhausting the resources of the affected trees. If
the trees be young or small at the time of attack their growth
is slowed up and they become misshapen, but their foliage re¬
mains green and their existence is not seriously threatened for
many years. The effect is of such a character that rusted
trees have been passed by without suspicion of the cause of
their condition.
Infection takes place in the needles of the current season.
Except for a very slight paling of the normal green there is no
indication of the rust for two years. It is only at the beginning
of the third year that the spermogonia mature, to be followed
about a month later by the peridermia. Even yet the affected
needles have lost little of their normal green aspect. From the
region of primary infection the mycelium invades the support¬
ing stem, then extends slowly both downwards and upwards
through the cortex of the latter, and thence out into the con¬
tiguous needles ; but the spread seems to involve not more than
a single internode in a season. Some of the needles attached
to an infected internode remain free from the rust, for the first
season at least. Those that are infected in this secondary way
behave like those primarily infected. Eventually they bear a
crop of spermogonia and peridermia. They usually drop dur¬
ing the following winter, but some continue to adhere and may
produce additional spermogonia and peridermia the following
season.
The mycelium in both stem and leaves develops large haus-
toria. These have been studied in detail by Miss L. M.
Hunter. Those in the stem are especially large and are char¬
acterized by remarkable, closely coiled, spiral forms.
94
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
The spermogonia are notable for their large size, their loca¬
tion under the epidermis, their enormous production of sper-
matia, and their long period of spore discharge. The dis¬
charge may continue for at least a month. It is because of
their large size that the aecial stage was called Peridermium
pycno grande by Bell. When the atmosphere is humid the
spermatia and spermatial fluid are extruded in great quantities.
I have many times seen the affected foliage wholly wetted by
the outflow and a drip falling from the leaves. This drip is
sweetish to the taste but not noticeably odorous.
M. polypodophila is not perennial on its fern host. But the
same result follows practically because of the fact that it
overwinters on the fronds, and abundant uredinial reproduc¬
tion occurs on these fronds in their second season. In this
way the new fronds are exposed to profuse inoculation with
uredospores, and infection freely follows. The rust propa¬
gates itself indefinitely on the same fern plant in this way. In
this respect, of course, M. polypodophila is like other species
of Milesia.
The peridium of the uredinium is unusual from the circum¬
stance that it is so slightly attached to the overlying epidermis.
The latter loosens and is readily pulled off without disturbance
to the uredinium or its peridium.
The teliospores are of strikingly large size, in keeping with
the large size of the host cells. They form extensive crusts,
the individual spores separated from one another only by the
anticlinal walls of the containing epidermal cells. The fre¬
quency with which they occur in the guard cells is worthy of
remark, as is also their absence from the upper epidermis.
The teliospores form in the spring at just about the time the
new foliage of Abies is breaking bud. Their cells germinate
without a resting stage, each one giving rise to a single four-
celled basidium.
Quite recently collections of this rust have been made in
New England (Clinton and Hunt, 26; Faull; Faull and Ches¬
ter), Pennsylvania (Thurston, 47), and as far south as Ten-
nesee (Whetzel et al.). The characters they manifest are
entirely typical of the species. Thus forty uredospores of the
Tennessee collection gave a measurement range of 14-23 x
45-62 n, with an average of 18 x 52 n; seventeen uredospores
1932]
FAULL, THE GENUS MILES I A
95
of a Maine collection measured 17-23 x 43-62 with an aver¬
age of 20 x 52 m. One other spore in the latter collection
measured 24 x 80 m-
That Bell (6), who had not yet seen the teliospores, referred
the uredinial stage to Uredinopsis is not surprising, because
the uredospores, though smooth, by their elongated and more
or less acuminate shape suggest that genus. In many ways
this is a remarkable Milesia in both of its host phases and in
its effect on its fir host. Plainly, however, it is not a Uredinop¬
sis, but in its essential characters falls into the genus Milesia.
27. Milesia laeviuscula (Dietel & Hoi way) Faull, n. comb.
II and III.
Uredo laeviuscula Dietel & Hoi way in Erythea, 2:127 (1894).
Thekopsora laeviuscula Dietel in Annal. Mycol. 1:416 (1903).
Hyalopsora laeviuscula Arthur in North Amer. Flora, 7:113
(1907).
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely aggregate on greenish to brownish areas of indefinite
extent, pustular, round, 0.2 to 0.5 mm. in diameter, covered by
yellowish brown epidermis which finally ruptures at a centrally
placed stomatic pore, or by an irregular slit; peridium hemi¬
spheric, firm, colorless, very strongly developed; peridial cells
isodiametrically to irregularly polygonal, 9 to 18 m across, with
hyaline walls 2 to 9 m thick; uredospores white, very short-
stalked, obovoid, ellipsoid or rarely subspherical, often quite
irregular in shape, 14-22 x 23-48 m, averaging about 17 x 33 m;
spore walls smooth, colorless, thin, about 1 m thick.
III. Telia on overwintered fronds, hypophyllous, on brown
areas of indefinite extent; teliospores within the epidermal
cells, occasionally in the guard cells, hyaline, rounded, or irreg¬
ular in outline and conforming to the shape of the containing
epidermal cell, often completely filling it, one- to many-celled,
up to forty or more, with vertical septa, a single pore in the
outer wall of each cell; the cells of the teliospores with thin,
smooth, colorless walls, irregularly polygonal except along
their free margins, 10-20 x 12-25 m; basidia 4-celled, about 7 m
thick and up to 55 m long; basidiospores spherical to ellipsoid,
7-8 x 7-11 m-
96
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Hosts and Distribution:
O and I. Unknown.
II and III. Polypodium calijornicum Kaulf., in California.
(M. laeviuscula forma typica).
Poly podium glycyrrhiza D. C. Eaton, in Oregon, Washing¬
ton, Alaska. (M. laeviuscula forma glycyrrhiza) .
Type Locality: Mt. Tamalpais, California, on Poly podium cali¬
jornicum. II and III.
Illustrations: Milesia laeviuscula typica Plate I, Figs. 2 a-d;
Text Fig. 1. — M. laeviuscula glycyrrhiza Plate I, Figs. 3 a-d; Text
Fig. 2.
Exsiccati: Barth. N. Am. Ured. 1505.
Specimens Examined. —
A. Milesia laeviuscula forma typica on Polypodium cali¬
jornicum.
California: Mt. Tamalpais, May 30, 1893, W. C. Blasdale (II,
III), telia on overwintered fronds; type. — Mt. Tamalpais,
June 15, 1894, W. C. Blasdale, Barth. N. Am. Ured. 1505 (II,
III), telia on overwintered fronds. — Monrovia, Mar. 11,
1922, M. S. Clemens, Arthur Herb. 4565 (II). — Monrovia,
April 29, 1922, M. S. Clemens, communicated by E. Bethel,
Arthur Herb. (II); on the packet Mr. Bethel asks about the
“white fungus”; the specimens show heaps or tendrils of
white, extruded spores. — Lake Phoenix, Marin Co., April
1928, overwintered fronds, H. E. Parks, Crypt. Herb. Univ.
of Calif. (II, III). — Mt. Tamalpais, April 3, 1932, Lee
Bonar, overwintered fronds (II, III); type locality.
B. Milesia laeviuscula forma glycyrrhiza on Poly podium
glycyrrhiza.
Washington: Bingen, May 12, 1904, Wilhelm N. Suksdorf (973),
Myc. Herb. Univ. of Wash. 738 (II). — Klickatat R., near
Mt. Paddo, May 8, 1886, overwintered fronds, W. N. Suks¬
dorf, Herb. N. Y. Bot. Gard. (II, III). — Klickatat R., May
8, 1886, W. N. Suksdorf, Myc. Herb. Univ. of Wash. 563
(II).
Oregon: Rhododendron, July 16, 1929, G. D. Darker (2800),
Arnold Arb. Path. Herb. 295 (II, III), telia on overwintered
fronds. — Rhododendron, July 20, 1929, G. D. Darker (2838),
Arnold Arb. Path. Herb. 333 (II, III), telia on overwintered
fronds. — Ashland, Aug. 1, 1929, G. D. Darker (2895),
Arnold Arb. Path. Herb. 390 (II, III), telia on overwintered
fronds. — Corvallis, undated, H. S. Jackson and F. D. B.,
Herb. Ore. Agr. Coll. 1104 (II). — Mt. Jefferson, July 27,
1907, E. R. Lake, Herb. Ore. Agr. Coll. 2508 (II, III), telia
on overwintered fronds. — Corvallis, Mar. 25, 1915, G. B.
Posey, Herb. Ore. Agr. Coll. 2626 (II). — Bridal Veil, May
18, 1915, H. S. Jackson, Herb. Ore. Agr. Coll. 3025 (II). —
1932]
FAULL, THE GENUS MILESIA
97
Hood River, May 16, 1915, II. S. Jackson, Herb. Ore. Agr.
Coll. 3042 (II). — Rhododendron, Sept. 6, 1931, /. H. Fatdl
(II).
Alaska: Sitka, June 22, 1916, P. J. Anderson (309), Herb. Alaska
Agr. Expmt. Sta. (II, III), telia on overwintered fronds. —
Juneau, June 3, 1917, overwintered fronds, P. J. Anderson,
Herb. Alaska Agr. Expmt. Sta. (II, III).
Milesia laeviuscula is based on specimens of rusted Poly¬
podium californicum collected by Professor W. C. Blasdale,
May 30, 1893, on Mount Tamalpais, near San Francisco, Cali¬
fornia. Dietel (10) studied this material and published a brief
account of the rust in 1894 under the name Uredo laeviuscula
Dietel and Holway, n. sp. His account made reference to the
uredinial stage only; but later, knowing better what to look
for, he discovered the telia in the same collections, and in 1903
he enlarged his description so as to include the telial stage
(11). His observations on the teliospores were comprehen¬
sive. He noted that they occurred on overwintered fronds in
lesions of indefinite extent, that they were many-celled, that
they usually filled the epidermal cells occupied, that some of
them were housed in guard cells, and that the basidia origi¬
nated one from each telial cell leaving tiny apertures in the
walls through which they passed. Having located the telio¬
spores the next step was to refer the rust to its proper genus.
For this purpose, however, one essential character had been
overlooked, namely, the color of the rust. Dietel hesitated
between two genera, both yellow-spored, namely, Melamp-
sorella and Thecopsora, but gave preference to the latter on
the grounds of septation of the teliospores. Presently Arthur
(5) transferred the species to Hyalopsora, likewise a yellow-
spored genus. But in reality the rust is white, quite without
pigment, and in this feature as well as in morphological char¬
acters, it unquestionably belongs in the genus Milesia.
That the uredospores are white had not gone unnoticed,
however. Searching through the Arthur Herbarium I found
two packets of rusted Polypodium californicum from southern
California on one of which the sender, Mr. E. Bethel, had pen¬
cilled an inquiry as to the identity of the “white” fungus on
his fern material. During the summer of 1931 it was my good
fortune to meet Mr. H. E. Parks of the University of Cali¬
fornia Herbarium, a collector of this rust on P. californicum,
98
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
who informed me that he was familiar with the fact that the
uredospores of M. laeviuscula are white as seen in extruded
masses on affected fronds in the field. I made a special effort
to see the rust for myself on the host in situ. Early in Sep¬
tember Professor Blasdale kindly took me to the exact region
from which he had gathered the type material; we found plenty
of the ferns but owing to a long continued drought the fronds
were entirely dried up. Professor Lee Bonar who accom¬
panied us on our trip, mindful of my quest, re-visited Mt.
Tamalpais on April 3, 1932. He found both the uredinial and
the telial stages of the rust on P. californicum at the same spot
1 2
Text Fig. 1. Surface view of part of the peridium from uredinium of Mile-
sia laeviuscula forma typica on Polypodium californicum.
Text Fig. 2. Surface view of part of the peridium from uredinium of Mile-
sia laeviuscula forma glycyrrhiza on Polypodium glycyrrhiza.
from which the type material had been collected in 1893, and
immediately forwarded specimens in fresh condition by air
mail. They arrived in excellent condition and were kept under
observation for many days in a moist chamber. I am able
now to state with certainty that the uredospores are snowy
white, without a trace of color. Search for the living rust on
P. glycyrrhiza in September, 1931, was soon rewarded by find-
1932]
FAULL, THE GENUS MILESIA
99
ing ample material at Rhododendron, near the foot of Mt.
Hood, in Oregon. Here and there on lesions on the living
fronds were to be seen uredinia with masses or tendrils of pure
white spores emerging from their apices.
This now leads to an inquiry into the exact identity of the
Milesia on Polypodium glycyrrhiza, interpreted to be M . laevi-
uscula, a rust distributed on this host from Alaska to Oregon.
Is it in reality M. laeviuscula , or is it a distinct but closely re¬
lated species? Heretofore no question has been raised with
regard to this point; the rusts on the two specific hosts have
been regarded as one and the same. There is a slight, though
perhaps not constant difference, however. I have found that
the uredo-peridial cells of the rust as found on P. glycyrrhiza
have walls that are rarely more than 2 \a thick, while those
from the rust on P. calif ornicum are thicker, ranging from 2 to
9 m (Text Figs. 1 and 2). It is possible that when all stages
are known and cross host inoculations have been made we
shall find that we are dealing with two distinct species or vari¬
eties; but meanwhile it seems better to dispose of them as the
one species, represented by the form M. laeviuscula typica
Faull on P. calif ornicum and M. laeviuscula glycyrrhiza Faull
form, nov., on P. glycyrrhiza.
The uredospores of M. laeviuscula , as is true of so many
species of Milesia, exhibit considerable variation, even on the
same specific host. Some details on this feature are recorded
in the accompanying table. In each case the data were se¬
cured from measurements on 40 spores taken at random from
individual uredinia.
Location
TABLE 7
Date Range
On Polypodium californicum
Average
Mt. Tamalpais, Calif, (type)
May 30, 1893
17-21
X
25-41 p,
17
x 34 p
Monrovia, Calif.
Apr. 29, 1922
14-20
X
28-45
17
x 35
L. Phoenix, Marin Co., Calif.
April 1928
14-20
X
25-40
17
x 33
B. On Poly podium glycyrrhiza
Rhododendron, Ore.
July 16, 1929
14-21
X
23-37
16
x 30
Rhododendron, Ore.
Sept. 6, 1931
14-20
X
25-42
16
x 32
Klickitat R., Wash.
May 8, 1886
14-20
X
28-48
15
x 38
Bingen, Wash.
May 12, 1904
14-20
X
23-40
15
x 32
Sitka, Alaska
June 22, 1916
15-22
X
28-40
17
x 36
Juneau, Alaska
June 3, 1917
15-20
X
25-34
17
x 32
100
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
A rust with smooth uredospores on Polystichum munitum
was referred by Jackson (28) to M. laeviuscula under the
name Hyalopsora laeviuscula ; but for various reasons this was
a misinterpretation. I have examined the rust in question; it
answers to M. vogesiaca , a rust quite common in Oregon and
quite likely elsewhere in western America, but not previously
reported from the New World.
The life histories of the two forms of M. laeviuscula remain
to be worked out. Such could probably be done quite easily,
for both forms are accessible, teliospores abound on over¬
wintered fronds and they germinate freely. It may be antici¬
pated that Abies is the alternate host.
I am deeply indebted to Professor Blasdale for a portion of
the type material on Polypodium californicum, and to Pro¬
fessor Lee Bonar for a collection of the rust in a living condi¬
tion from the type locality.
X. SPECIES ON POLYSTICHUM
28. Milesia exigua Faull, n. comb. O, I, II and III.
Milesina exigua Faull in Jour. Arnold Arb. 12:218-9 (1931).
II.
Milesina vogesiaca S. Kamei in Trans. Sapporo Nat. His.
Soc. 11:141-147 (1930), not Sydow. O, I, II and III.
O. Spermogonia on needles of current season, amphige-
nous, mostly hypophyllous, inconspicuous, colorless, im¬
mersed, inverted hemispherical in sectional view, 120-175 p
broad by 110-160 p deep; spermatiophores unbranched, sep¬
tate; spermatia hyaline, unicellular, smooth, narrowly ellip¬
soid, 1.5-2 x 5-7 p.
I. Aecia hypophyllous on needles of current season, in two
rows, one on each side of midrib, white, cylindrical, 0.25 mm.
in diameter by 0.5 mm. high; peridium colorless, delicate, rup¬
turing at the apex; peridial cells polygonal, elongated verti¬
cally, overlapping, in a single layer, colorless, 14-22 x 26-33 p,
inner wall thick, closely verrucose, outer wall thin, smooth;
aeciospores globose to ellipsoid, 14-22 x 19-25 p, mostly 18 x
21 p, wall thin, white, verrucose.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite
extent, pustular, 0. 1-0.2 mm. in diameter, rupturing at a cen-
1932]
FAULL, THE GENUS MILESIA
101
trally placed stomatic pore in the overlying brownish epi¬
dermis; peridium hemispheric, hyaline, delicate; peridial cells
isodiametrically to irregularly polygonal, 8-12 x 8-15 p; walls
of peridial cells hyaline, 0.5 to 1.0 p thick; uredospores color¬
less, smooth, thin-walled (0.5- 1.0 p), short-stalked, obovoid to
ellipsoid or subspherical, 14-17 x 18-29 p, averaging about
15 x 24 p.
III. Telia on fronds of current season, mostly hypophyl-
lous, on indefinite, extensive, brown areas; teliospores within
the epidermal cells, even the guard cells, hyaline, rounded, or
irregular in outline and conforming to the shape of the con¬
taining epidermal cell, usually completely filling it; few- to
many-celled, up to thirty or more, with anticlinal septa; the
cells of the teliospores with thin, smooth, colorless walls, and
a single germ pore, irregularly polygonal except along their
free margins, 9-20 x 9-23 p.
Hosts and Distribution:
O and I. Abies Mayriana Miyabe and Kudo, A. firma Sieb.
and Zucc., A. sachalinensis Mast., in Japan. Cultured from
Polystichum Braunii (29).
II. Polystichum Braunii (Spenner) Fee, in Poland, Japan.
Polystichum tripteron (Kunze) Pr., in Japan.
Polystichum japonicum (Franch. et Sav.) Diels, in Japan.
Polystichum aculeatum (L.) Roth. var. retrorso-paleaceum
Kodama, in Japan.
III. Polystichum Braunii (Spenner) Fee, in Japan.
Polystichum tripteron (Kunze) Pr., in Japan.
Polystichum japonicum (Franch. et Sav.) Diels, in Japan.
Polystichum aculeatum (L.) Roth. var. retrorso-paleaceum
Kodama, in Japan.
Type Locality: Kniazdwor, Kolomea, Poland, on Polystichum
Braunii. II.
Illustrations: Plate V, Figs. 19 a-d. — S. Kamei in Trans. Sap¬
poro Nat. Hist. Soc. 11:143 and 145, Figs. 1-3 (1930).
Specimens Examined. —
A. On Abies firma.
Japan: Sapporo, a culture by S. Kamei, inoculation from Poly¬
stichum Braunii made June 17, 1926, peridermia appeared
July 14, 1926 (O, I).
B. On Polystichum Braunii.
Poland: Kniazdwor, Kolomea, Aug. 1913, A. Wroblewski (II);
type. — Olszanica, Lesko, Oct. 18, 1917, A. Wroblewski (II).
Japan: Nopporo Forest, near Sapporo, April 20, 1930, S. Kamei
102
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
(II). — Mount Moiwa, near Sapporo, Sept. 20, 1930, S.
Kamei (II, III).
C. On Poly stic hum tripteron.
Japan: Manisan, near Tottori, May 3, 1929, Naohide Hiratsuka,
overwintered fronds (II, III).
D. On Poly stic hum japonicum.
Japan: Mt. Daisen, Prov. Hoki, Nov. 17, 1929, Naohide Hiratsuka
(II, III).
E. On P olystichum aculeatum var. retrorso-paleaceum.
Japan: Mt. Daisen, Prov. Hoki, Nov. 9, 1930, Naohide Hiratsuka
(II, III).
The specimens from which Milesia exigua was described
were communicated by A. Wroblewski from Poland; they
occurred on Polystichum Braunii , and bore uredinia only. I
have recently received from Senji Kamei uredinial and telial
material on the same host from Japan, and spermogonial and
aeeial material on Abies firma, obtained by culturing from
teliospores on P. Braunii.
There has also come to my hand through the kindness of
Professor Hiratsuka three packets of rusts which closely resem¬
ble M. exigua , one each on the respective hosts Polystichum
tripteron, P. japonicum and P. aculeatum var. retrorso-palea¬
ceum. All bear teliospores as well as uredospores. As to the
first Professor Hiratsuka, who was probably not yet familiar
with M. exigua, interrogatively suggested the specific name
Milesina Polystichi-tripteri, n. sp. Its uredinia are certainly
close to those of M. exigua. Forty uredospores from this
material measured 12-17 x 22-31 p, with an average of 14 x
26 p; these spores are smooth, their walls are less than 1 p
thick, and the walls of the peridial cells measure 0.5 to 1.0 p
in thickness. On the basis of uredinia and uredospores I can¬
not separate this rust from M. exigua. Teliospores are pres¬
ent and they are on overwintered fronds, and in all respects
like those of M. exigua. The remaining organs, spermogonia
and aecia, are not yet known; the former might be morpho¬
logically different from those of M. exigua. So, too, the rusts
may be host restricted. For the present, however, the rust in
question on P. tripteron should in my opinion be diagnosed as
M. exigua, and with this conclusion Professor Hiratsuka is
now in agreement (51).
Concerning the rusts on the collections of Polystichum
1932]
FAULL, THE GENUS MILESIA
103
japonicum and P. aculeatum var. retrorso-paleaceum referred
to, I have reached the same conclusion; so far as their known
morphological characters are concerned they cannot be sepa¬
rated from M. exigua. We are unable, of course, to speak of
their spermogonia and aeoia because they remain to be dem¬
onstrated; nor is there any information on host restrictions.
Morphologically like typical M. exigua these particular rusts
also produce their teliospores in the fall on fronds of the cur¬
rent season just as Kamei has shown to be true of M. exigua
on P. Braunii in Japan.
We owe our knowledge of the life history of this rust to
Kamei (29). I am under deep obligations to this ardent in¬
vestigator for the opportunity of studying all stages of the rust
with which he worked. My observations on his materials co¬
incide closely with those recorded by him, except as to the
uredospores and the number of cells in the teliospores. The
uredospores I find to be smooth in wet mounts, and to measure
11-16.5 x 17-32 m; an average of 50 spores gave measures of
13.5 x 23.5 m- The spores are consistently much smaller than
those of M. vogesiaca, to which species he referred the rust.
The teliospores are abundant in both the lower and the upper
epidermes, and frequent in the guard cells; they are few- to
many-celled (up to 25 or more). On the other hosts in which
I found teliospores they were observed in the lower epidermis
only and not in the guard cells.
29. Milesia vogesiaca (Syd.) Faull, n. comb. II and III.
Milesina vogesiaca Sydow in Annal. Mycol. 8:491 (1910).
II and III (?).
Uredo vogesiaca (Syd.) Saccardo & Trotter in Syll. Fung.
21:812 (1912).
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brown areas of indefinite ex¬
tent, pustular, round to slightly elongate, 0.1 -0.3 mm. in
length, covered by a light brownish discolored epidermis with
centrally placed stomatic pore; peridium hemispheric, deli¬
cate; peridial cells hyaline, isodiametrically to irregularly
polygonal, 8-15 m across; walls of peridial cells hyaline, 0.5-1. 5
M in thickness; uredospores colorless, white in mass, numerous,
104
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
produced singly, short-stalked, obovoid or ellipsoid, 14-23 x
29-44 |j, on an average about 18 x 36 jj; walls of spores thin,
less than 1 m thick, smooth.
III. Telia hypophyllous and occasionally epiphyllous, on
indefinite extensive brown areas; teliospores within the epi¬
dermal cells, sometimes within even the guard cells, hyaline,
rounded, or irregular in outline and conforming to the shape
of the containing epidermal cell, usually completely filling it,
one- to fifty-celled, with vertical septa, a single pore in the
outer wall of each cell; the cells of the teliospores with thin,
smooth, colorless walls, irregularly polygonal except along
their free margins, 8-14 x 9-17 m-
Teleutosporae intra cellulas epidermidis evolutae, hypo-
phyllae, verticaliter septatae, pluricellulares (1-50 celluli),
leves, hyalinae; cellulae teleutosporarum 8-14 x 9-17 m-
Hab. in foliis Polystichi lobati in Polonia et Helvetia; P.
aculeati in Gallia; P. muniti in Oregon; P. varii in Japonia.
Hosts and Distribution:
O and I. Unknown.
II. Polystickum lobatum (Hudson) Presl, in France, Swit¬
zerland, Poland.
Polystickum Lonchitis (L.) Roth, in Switzerland.
Polystickum munitum (Kaulf.) Presl, in Oregon (U. S.)
Polystickum aculeatum (L.) Roth, in Algeria, France,
Czechoslovakia.
Polystickum varium (L.) Presl, in Japan.
III. Polystickum lobatum (Hudson) Presl, in Poland,
Switzerland.
Polystickum aculeatum (L.) Roth, in France.
Polystickum munitum (Kaulf.) Presl, in Oregon.
Polystickum varium (L.) Presl, in Japan.
Type Locality: Hohneck, Central Vosges, France, on Poly¬
stickum lobatum. II.
Illustrations: Plate V, Figs. 17 a-d; Plate IX, Fig. 34.
Exsiccati: Syd. Myc. Germ. 878. — Syd. Ured. 2345, 2741,
2742. — Maire Myc. Bor.-Afric. 287.
Specimens Examined. —
A. On Polystickum lobatum.
France: Hohneck, Central Vosges, July 12, 1910, H. Sydow, Syd.
Myc. Germ. 878 (II); type locality. — Hohneck, Vosges, July
16, 1910, H. Sydow, Syd. Ured. 2345, labelled Milesina voge-
siaca Syd. nov. spec. (II); type locality.
Switzerland: Crete de la Chaille, Neuchatel, June 22, 1925, over¬
wintered fronds, E. Mayor (II, III).
1932]
FAULL, THE GENUS MILESIA
105
Poland: Cetnerowka, Lwow, May 22, 1917, overwintered fronds,
A. Wroblewski (II, III).— Ojcow, July 28, 1925, Wanda
Konopacka , Herb. W. Siemaszko (II).
B. On Polystichum Lonchitis.
Switzerland: Leysin, Vaud, Sept. 26, 1921, E. Mayor (II).
C. On Polystichum munitum.
Oregon: Mary’s Peak, Corvallis, Apr. 23, 1915, G. B. Posey, Herb.
Ore. Agr. Coll. 3041 (II). — Shell Rock Lake, Clackamas Co.,
July 8, 1929, overwintered fronds, G. D. Darker (2674),
Arnold Arb. Path. Herb. 171 (II, III). — Oregon Caves, Jose¬
phine Co., Aug. 10, 1929, overwintered fronds, G. D. Darker,
Arnold Arb. Path. Herb. 497 (II, III). — Lake Mt. Trail,
Oregon Caves, Aug. 12, 1929, G. D. Darker (3039) (some
M. Polystichi in same packet), Arnold Arb. Path. Herb. 534,
overwintered fronds (II, III). — Camp Creek, Rhododendron,
Sept. 6, 1931, /. H. Fault (II). — Rhododendron, Sept. 6,
1931, J. H. Fault (II).
D. On Polystichum varium.
Japan: Tottori, Mar. 17, 1930, overwintered fronds, Naohide
Hiratsuka (II, III). ( Milesia vogesiaca forma nipponensis) .
E. On Polystichum aculeatum.
Algeria: Djurdjura, May 25, 1915, R. Maire, Maire Mye. Bor.-
Afric. 287 (II).
France: Foret des Fangs, Axat, Aude, Apr. 30, 1932, G. D. Darker
and G. Fenwick-Owen (II, III).
Czechoslovakia: Kralovany, Slovakia, July 26, 1932, G. D. Dar¬
ker (II).
Milesia vogesiaca under the name Milesina vogesiaca was
originally described (43) from uredinial material collected by
H. Sydow on fronds of Polystichum lobatum in the central
Vosges. At the time it was the only known species of Milesia
with smooth uredospores, and so was regarded as more or less
anomalous. There was uncertainty as to the presence of telia
in the type collection; as the gathering was made in July it
may well have included overwintered fronds with some telio-
spores. Collections of telial as well as uredinial material on
the same host, here recorded for the first time, were later made
by A. Wroblewski in Poland and Eugene Mayor in Switzer¬
land, in both instances on overwintered fronds; it is from these
that I have described the telia as above. Besides P. lobatum
it now appears that we should add as hosts for this rust P.
munitum, P. Lonchitis, P. varium and P. aculeatum.
On examining several collections of rusted Polystichum
106
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
munitum from Oregon made by Dr. G. D. Darker in 1929, I
found three collections, all on overwintered fronds, which car¬
ried a white rust with smooth uredospores and septate, intra-
epidermal teliospores. One collection (Path. Herb. Arnold
Arb. 171) was made July 8, 1929, at Shell Rock Lake, Oregon,
elev. 4000 ft.; a second collection (idem 497) was made Aug.
10, 1929, at Oregon Caves, Oregon, elev. 4300 ft.; and the
third (idem 534) on Aug. 12, 1929, also at Oregon Caves. To
these I have added two collections of my own made in Oregon
in 1931. Looking through the literature I discovered a record
by Jackson (28) of a rust on Polystichum munitum from Ore¬
gon, made by G. B. Posey, April 23, 1915 (Herb. Ore. Agr.
Coll. 3041), which was diagnosed by the former as Hyalopsora
laeviuscula (Diet.) Arthur. Through the courtesy of Pro¬
fessor H. P. Barss I have been enabled to re-examine this
material, and have found that it is the same as in Darkens
collections referred to above, and not as determined by Jack-
son.
What then is the white-spored rust on Polystichum munitum
collected by Posey, by Darker and by Faull? Plainly enough
it is a Milesia. Posey’s material was collected too early in the
season to show teliospores, but they are abundant in Darker’s
collections; so the genus is fixed. It may be necessary to
know the aecial stage before a final specific determination can
be made, because experience has shown that the spermogonia
in Milesia are of diagnostic value. It would be desirable, too,
to test by cultures on other allied hosts. However, on the basis
of morphological characters of uredospores and teliospores,
especially the former, there is nothing that would enable one
to distinguish it from M. vogesiaca. Therefore, I have tenta¬
tively referred it to this species.
There yet remain to be mentioned three other collections —
a Milesia on Polystichum Lonchitis in Switzerland, a uredinial
collection of which was sent to me by Dr. Eugene Mayor,
gathered near Leysin (Alpes vaudoises), Canton de Vaud, on
Sept. 26, 1921; a Milesia on P. varium from Japan; and a
Milesia on P. aculeatum from Algeria and from the south of
France. A careful study of the uredinia and uredospores of
the first shows that they present such a close morphological
resemblance to those of M. vogesiaca on P. lobatum that I
1932]
FAULL, THE GENUS MILESIA
107
feel justified in also referring this rust tentatively to M. voge-
siaca. The only constant difference noted is the slightly
thicker walls of the uredinial peridial cells. A final determina¬
tion must await comparative studies of spermogonia and peri-
dermia, and cross inoculations on the respective fern hosts.
A similar disposition for similar reasons is made of the rusts
on P. varium and P. aculeatum.
A comparison of uredospores studied by me from these vari¬
ous hosts is presented in Table 8. The averages given are of
40 measured spores each, taken as they came in the mounts.
The bracketed numbers in the second column serve to identify
the packets as filed in my own collection.
TABLE 8
Host Collection
P. lob alum Syd. Myc. Germ. 878
P. lobatum Wroblewski, Poland
(9884)
P. lobatum Mayor, Switz. (9882)
P. Lonchitis Mayor, Switz. (9883)
P. munitum Herb. Univ. Oregon
3041
P. munitum Herb. Arn. Arb. 171
(Oregon)
P. varium Hiratsuka (10374)
P. aculeatum Mayor, Myc. Bor.-Afric.
287
Average size
of uredospores
17.8 x 32.4 \i
19.3 x 36.9
19.2 x 32.4
19.2 x 35.7
18.7 x 37.3
17.9 x 36.4
18.3 x 36.0
18.0 x 36.0
Range in size
of uredospores
14.2-22.7 x 25.6-42.6 n
17.0-22.7 x 28.4-46.9
17.0-22.7 x 28.4-38.3
14.2-22.7 x 28.4-45.4
14.2-22.7 x 27.0-45.4
14.5- 23.2 x 29.0-43.5
15.6- 21.3 x 28.4-45.4
16.4-24.6 x 27.4-41.1
As yet we have no knowledge of the aecial phase of M. voge-
siaca. The rust whose aeoia were described by Kamei (29)
under that name answers to M. exigua. We may reasonably
anticipate, however, that successful cultures could be carried
out on Abies with the teliospores of M. vogesiaca from its vari¬
ous telial hosts.
In conclusion a brief comment should be made with refer¬
ence to the period of teliospore formation. So far teliospores
have been found on the hosts Polystichum lobatum and P.
aculeatum in Europe, P. munitum in North America, and P.
varium in Japan. I have examined sufficient material of the
first three to lead to the conclusion that the teliospores develop
in ordinary course on overwintered fronds, and not at the close
of the preceding season. The one lot of the telial stage on P.
108
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
varium I have seen was gathered about the middle of March
on overwintered fronds; collected at a date so early in the
spring it seems quite likely that the teliospores present were
actually formed in the preceding late fall on fronds of the same
year. If this be true, as it would seem to be, then two habits
are represented by the rust recognized here as M. vogesiaca,
namely, teliospore formation in the fall on leaves of the current
season and teliospore formation in the early part of the fol¬
lowing growing season on overwintered fronds. The latter
may be considered as typical for the species, while tentatively
the former may be characterized as the habit of a form of the
species. I shall designate this form M. vogesiaca forma
nipponensis Faull, f. nov. What culture studies of M. vogesiaca
on its various hosts as denoted above would show we do
not know. Quite possibly they would show definite host re¬
striction. Nor do we know anything of the aeeial stages of
the rust as derived from its respective hosts. They might very
well show morphological differences, especially as to their
spermogonia. Considering the geographic and host distribu¬
tion, the species as defined in this paper, may be palingenetic
in its composition; but as yet there is no evidence indicative
as to whether M. vogesiaca is a natural specific unit, an aggre¬
gation of biological strains, or a group of two or more con¬
vergent species now morphologically alike.
30. Milesia Polystichi Wineland in H. S. Jackson in Mem.
Brookl. Bot. Gard. 1:214 (1918). (II). II and III.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite
extent, pustular, round, 0.2 -0.3 mm. in diameter, covered by
a brownish discolored epidermis which finally ruptures at a
centrally placed stomatic pore; peridium hemispheric, firm;
peridial cells hyaline, isodiametrically to irregularly polygonal,
8-16 m across; walls of peridial cells hyaline to brown, 1-3 m
thick; uredospores colorless, white in mass, numerous, pro¬
duced singly, short-stalked, obovoid or ellipsoid, 15-20 x 20-
35 m, averaging about 17 x 28 p; walls of spores thick, 1.5-
2.5 p, strongly echinulate.
III. Telia on overwintered fronds, hypophyllous and occa-
1932]
FAULL, THE GENUS MILES I A
109
sionally epiphyllous, on indefinite, extensive brown areas, at
times involving entire pinnae ; teliospores within the epidermal
cells, sometimes within the guard cells, hyaline, rounded, or
irregular in outline and conforming to the shape of the con¬
taining epidermal cell, often completely filling it, 1- to many-
celled, with vertical septa, a single pore in the outer wall of
each cell; the cells of the teliospores with thin, smooth, color¬
less walls, irregularly polygonal except along their free mar¬
gins, 8-16 x 11-23 m-
Teleutosporae intra cellulas epidermidis evolutae, amphige-
nae, praecipue hypophyllae, verticaliter septatae, pluricellu-
lares, leves, hyalinae; cellulae teleutosporarum 8-16 x 11-23 q.
Hab. in foliis Polystichi muniti in Oregon.
Hosts and Distribution:
O and I. Unknown.
II. Polys tic hum munitum (Kaulf.) Presl, in Oregon, Cali¬
fornia, Idaho, Montana, Washington.
III. Polystichum munitum (Kaulf.) Presl, in Oregon. The
telial stage is reported for the first time and from Oregon only;
but it probably occurs throughout the range of the rust.
Type Locality: Grant’s Pass, Josephine Co., Oregon, on Poly¬
stichum munitum. II.
Illustrations: Plate V. Figs. 18 a-d.
Specimens Examined, all on Polystichum munitum. —
Oregon: Grant’s Pass, Sept. 5, 1916, /. R. Weir (260) (II); part
of type. — Zig Zag R., Government Camp, July 15, 1929,
G. D. Darker (2788) and L. N. Goodding, Arnold Arb. Path.
Herb. 283 (II, III), overwintered fronds. — Lake Creek, Ore¬
gon Caves, Aug. 11, 1929, G. D. Darker (3021), Arnold Arb.
Path. Herb. 516 (II, III), overwintered fronds. — Lake Mt.
Trail, Oregon Caves, Aug. 12, 1929, G. D. Darker (3039),
Arnold Arb. Path. Herb. 534 (II, III), overwintered fronds.
— Zig Zag R., Rhododendron, July 20, 1929, G. D. Darker
(2841), Arnold Arb. Path. Herb. 336 (II, III), overwintered
fronds. — Government Camp, July 22, 1929, G. D. Darker
(2855), Arnold Arb. Path. Herb. 350 (II). — Zig Zag Mt.
Trail, Rhododendron, July 9, 1929, G. D. Darker (2686),
Arnold Arb. Path. Herb. 183 (II).
California: Santa Cruz, Apr. 19, 1920, E. Bethel (II). — Requa,
Aug. 20, 1929, G. D. Darker (3132), Arnold Arb. Path.
Herb. 627 (II).— Requa, Sept. 1918, /. R. Weir (12679)
(II).
Washington: Langley, Aug. 1923, J. M. Grant, Herb. Bur. PI.
Ind., U. S. Dept, of Agr., Washington, D. C. (II). — Langley,
110
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
1920, J. M. Grant, Farlow Herb. (II). — Forks, Olympic
Peninsula, Sept. 1, 1931, J. H. Fault (II). — Sol Due Springs,
Chalum Co., July 14, 1921, /. R. Weir, Herb. Bur. PI. Ind.,
U. S. Dept, of Agr., Washington, D. C. 19903 (II). — Chelan
Lake, Aug. 25, 1916, J. R. Weir, Herb. Bur. PI. Ind., U. S.
Dept, of Agr., Washington, D. C. 7534 (II).
Montana: Troy, Aug. 1913, /. R. Weir, Herb. Bur. PI. Ind., U. S.
Dept, of Agr., Washington, D. C. 12765 (II).
Idaho: Kooskia, Sept. 1915, J. R. Weir, Herb. Bur. PI. Ind., U. S.
Dept, of Agr., Washington, D. C. 12889 (II).
The uredinia of Milesia Polystichi are abundant and con¬
spicuous on the lesions, particularly on the overwintered
fronds. Their peridia have been described as being marked
by radially elongate cells. Vertical sections show that not
only is this the case, but also that there may be several in the
same radius. As explained elsewhere, these are primordial
spore mother columns from which the peridial cells proper
have not yet been organized. Incidentally it may be noted
that peridia are so firm that they can in part be freed quite
easily in wet mounts from the overlying epidermis.
The telia, as is generally true of species of Milesia, develop
in the lesions on overwintered fronds.
As yet the aecial stage of this rust has not been established
by cultures. It would apparently be a simple matter to do so,
and it is anticipated that Abies will prove to be the alternate
host. Where Firs grow near rusted Poly Stic hum munitum
they frequently carry large crops of white peridermia, and the
association is such as to lead one to reasonably suspect that
these peridermia constitute the aecial phase of the fern rust.
But the rust on Polystichum munitum may either be M. Poly¬
stichi, or the frequent, smooth-spored rust hitherto unrecog¬
nized in America, diagnosed in this paper as M. vogesiaca.
Peridermium rugosum Jackson has been suggested as the hap¬
loid stage of M. Polystichi, but without proof (25). Perhaps
it is; on the other hand perhaps it is M. vogesiaca, or even
some other one of the five known western American species of
Milesia; before reaching a final decision controlled cultures
will be necessary. Such cultures will be important, because
they will establish the life history of the species, and they may
determine the status of Peridermium rugosum.
Grove (22) listed a rust on Polystichum aculeatum in Eng-
1932]
FAULL, THE GENUS MILESIA
111
land under the name Milesina Polys tic hi (Wineland) Grove;
but the rust in question is not the Milesia Polystichi of Wine-
land. It is described in this paper under the name Milesia
Whitei.
31. Milesia Whitei Faull, sp. nov. II et III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide teeti, sparsi vel aggregate pustulati, rotundati,
0.15-0.3 mm. diam., peridio ex cellulis hyalinis, peridio interne
ex cellulis radiatim elongatis superne irregulariter polygonali-
bus composite cincti; cellulae peridii imbricatae, 8-15 p diam.,
pariete 0. 7-0.9 p crasso. Uredosporae obovatae vel ellip-
soideae, remote et subtiliter echinulatae, hyalinae, 17-22 x
22-40 p, plus minusve circa 19 x 30 p, episporio 0. 5-0.9 p
crasso. Teleutosporae in hypophyllo intra cellulas epidermi-
dis evolutae, verticaliter septatae, pluricellulares, leves, hya-
linae; cellulae teleutosporarum 6-15 x 8-20 p.
Hab. in foliis Polystichi aculeati in Yugoslavia, Caucasia et
Britannia.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, subepidermal, scattered or
loosely grouped on greenish to brownish areas of indefinite
extent, inconspicuous, pustular, round, 0.15 to 0.3 mm. in
diameter, covered by a buff discolored epidermis which finally
ruptures at a central, stomatic pore; peridium very delicate,
hemispheric; peridial cells hyaline, small, isodiametrically to
irregularly polygonal, overlapping, more or less radially
elongate next to the base, 8-15 p across; walls of peridial cells
hyaline, less than 1 p thick; uredospores colorless, white in
mass, short-stalked, obovoid or ellipsoid, rarely subspherical,
17-22 x 22-40 p, averaging about 19 x 30 p; walls of spores
thin, less than 1 p thick, rather sparsely and finely echinulate.
III. Telia on overwintered fronds, hypophyllous, on in¬
definite, extensive, brown areas, at times involving entire pin¬
nae; teliospores within the epidermal cells, hyaline, rounded,
or irregular in outline and conforming to the shape of the con¬
taining epidermal cell, often completely filling it, one- to many-
celled, with anticlinal septa; the cells of the teliospores with
rather thin, smooth, colorless walls, irregularly polygonal ex¬
cept along their free margins, 6-15 x 8-20 p.
112
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Hosts and Distribution:
O and I. Unknown.
II. Polystichum aculeatum (L.) Roth, in Yugoslavia, Cau¬
casus, England.
III. Polystichum aculeatum (L.) Roth, in Yugoslavia.
Type Locality: Castel Nuovo, Cattaro Bay, Dalmatia, Yugo¬
slavia, on Polystichum aculeatum. II and III.
Illustrations: Plate V, Figs. 20 a-d.
Exsiccati: Jaap Fung. Sel. 719, under name Milesina Krieg-
eriana Magn.
Specimens Examined, all on Polystichum aculeatum. —
Yugoslavia: Castel Nuovo, April 25, 1914, O. Jaap , Jaap Fg. Sel.
719, overwintered fronds (II, III); type.
Caucasus: Suchum, Georgia, Nov. 1918, W. Siemaszko , Herb. Bur.
PI. Ind., U. S. Dept, of Agr., Washington, D. C., under name
Milesina Theissenii Siemaszko, n. sp. (inedit.) (II).
England: Tintern, April 1925, E. M. Wakefield, Herb. Hort. Bot.
Reg. Kew under name Milesina Polystichi (Wineland) Grove
(II).
Milesia Whitei appears to have been more or less of a puzzle
to uredinologists who have made reference to it. As noted
above three separate collections found in herbaria are desig¬
nated by as many different names. Siemaszko indicated that
it might be an undescribed species; he labeled his gathering
(without description) Milesina Theissenii, n. sp.; but he later
regarded it as M. Polystichi (Wineland) Grove, according to
his statement in a letter dated May 12, 1932. Incidentally
it is to be observed that he identified the fern host as Poly¬
stichum Braunii (Spenner) Fee, but comparisons made in the
Gray Herbarium of Harvard University show that the host is
the closely related species, P. aculeatum (L.) Roth.
Miss Wakefield’s collection from England bears the fungus
in abundance and in excellent condition. Grove erroneously
diagnosed this collection as Milesina Polystichi (Wineland)
Grove, n. comb., the published report of which appears to be
the sole foundation of statements in the literature to the effect
that Milesia Polystichi occurs in Europe. The rust in ques¬
tion is certainly not M. Polystichi, nor has the latter been
found anywhere except in the western part of North America.
M. Whitei is easily distinguished from M. Polystichi in
several characters. The uredospores of the former are thin-
walled and are minutely spinulose, the uredinial peridium is
1932]
FAULL, THE GENUS MILESIA
113
delicate and the peridial cell-walls are thin; the uredospores
of the latter are thick-walled and quite strongly spinulose, the
uredinial peridium is firm and the peridial cell-walls are rela¬
tively thick. It may be anticipated that the spermogonia
when known will be found to present other differences, and it
is improbable that the species are host interchangeable.
Fortunately Jaap’s specimens from Jugoslavia contain
teliospores. His material is fairly abundant and is rightly
chosen as the type. No doubt teliospores could readily be
located on overwintered fronds at the proper season elsewhere
throughout the wide range of M. Whitei. Quite likely, too,
Abies will prove to be the alternate host.
This species is dedicated to Dr. F. Buchanan White, former
editor of the Scottish Naturalist, and the founder of the genus
Milesia.
XI. SPECIES ON SCOLOPENDRIUM
32. Milesia Scolopendrii (Fuckel) Arthur in Bull. Tor. Bot.
Club, 51:52 (1924). (II). II and III.
Ascospora Scolopendrii Fuckel in Symb. Myc. II, 2nd
Nachtr. 19 (1873). II.
Aecidium Scolopendrii (Fuckel) Oudemans in Bot. Zeitung,
33:585 (1875). II.
Uredo (?) pteridinm F. B. White in Scot. Nat. 4:27 (1877-8).
Puccinia pteridum F. B. White in Scot. Nat. 4:163 (1877-8).
IL
Uredo Scolopendrii Schroeter in Cohn, Krypt.-Flora Schles.
Ill, 1:374 (1889). II.
Uredinopsis Scolopendrii (Fuckel) Rostrup in Bot. Tidsskr.
21:42 (1897). II.
Milesina Scolopendrii (Fuckel) Jaap in Jaap Fg. Sel. 571
(1912). II.
O and I. Spermogonia and aecia unknown.
II. Uredinia hypophyllous, pustular, scattered or loosely
aggregate, frequently in rows between the lateral veins and
parallel to them, on greenish to brownish areas of indefinite
extent, sometimes involving almost the entire frond, round,
0.1 to 0.3 mm. in diameter, covered by brownish epidermis,
which finally ruptures at a centrally placed stomatic pore;
peridium colorless, hemispheric, delicate; peridial cells elon¬
gate and radially oriented at the sides of the peridium contigu¬
ous with the base, irregularly polygonal, isodiametric to some-
114
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
what elongated in the upper part of the peridium, radially
overlapping, 7 to 17 m across, with walls up to 1 n thick;
uredospores colorless, stalked (with stalks up to 16 m long),
obovoid or ellipsoid, 14-23 x 28-57 p, averaging about 19 x 37
m; spore wall hyaline, 0.5 to 1.5 m thick, quite strongly and
rather sparsely echinulate.
III. Telia hypophyllous, occasionally also epiphyllous, on
indefinite, brown areas on overwintered fronds; teliospores
within the epidermal cells, sometimes within even the guard
cells, hyaline, rounded, or irregular in outline and conform¬
ing to the shape of the containing epidermal cell, often com¬
pletely filling it, 1- to 40-celled, with anticlinal septa, at times
several one- to few-celled spores in a single epidermal cell;
the cells of the teliospores with thin, smooth, colorless walls,
and a single germ pore, irregularly polygonal except along
their free margins, 7-15 x 8-25 m; basidiospores globular to
subglobular, 6. 0-7. 5 m in diameter.
Teleutosporae intra cellulas epidermidis evolutae, amphig-
enae, praecipue hypophyllae, verticaliter septatae, pluricellu-
lares (1-40 celluli), leves, hyalinae; cellulae teleutosporarum
7-15 x 8-25 m-
Hab. in foliis Scolopendrii vulgaris ( Phyllitidis Scolo-
pendrii ) in Helvetia, Britannia et Gallia.
Hosts and Distribution:
O and I. Unknown.
II. Scolopendrium vulgar e Smith (1793) [Phyllitis Scolo-
pendrium (L.) Newm.], in Switzerland, Germany, Austria,
France, Italy, Great Britain, Poland, Spain, Caucasus.
III. Scolopendrium vulgar e Smith, in Switzerland, France,
England, as reported in this paper for first time. Probably
throughout range of the rust on overwintered fronds.
Type Locality: “In der Class im Ct. Graubiinden” (Grisons),
Switzerland, on Scolopendrium vulgare. II.
Illustrations: Plate VI, Figs. 22 a-c. — F. B. White in Scot.
Nat. 4: PI. II, Fig. 6 (1877-8). — R. Gonzalez Fragoso in Flora
Iberica, Uredales, II, Fig. 138, p. 282 (1925). — (?) M. C. Cooke in
Fg. Brit., 2nd ed., no. 73, thought by White (49) (probably cor¬
rectly so) to be drawn from spores of Hyalopsora Polypodii (DC.)
Magn. though on a mount of M. Scolopendrii under name Uredo
filicum Desm.
Exsiccati: Jaap Fg. Sel. 571. — Roum. Fg. Sel. Gall. 5212, 7084,
under name Uredo Scolopendrii (Fuckel) Schroet. — Syd. Ured. 2399
1932]
FAULL, THE GENUS MILESIA
115
under name Uredo Scolopendrii (Fuckel) Schroet., 2596 — Vester-
gren Microm. Rar. Sel. 1703, under name Milesina Dieteliam. —
Vise Fg. Brit. 143, under name Uredo flicum Desm. — Raciborski
Myc. Polon. 186, under name Uredo Scolopendrii (Fuckel) Rost.
(!)• — Cooke Fg. Brit. 2nd ed. 73, under name Uredo fHicum Desm.
Specimens Examined, all on Scolopendrium vulgare. —
Switzerland: In der Cluss im Ct. Graubiinden, late summer, L.
Fuckel, Herb. Barbey-Boissier, Univ. Geneve (II); type. —
Treymont, Neuchatel, June 12, 1920, E. Mayor, overwintered
fronds (II, III and basidia); type for teliospores. — Creux-
du-Vau, Jura neuchatelois, Oct. 19, 1913, E. Mayor, Syd.
Ured. 2596, also Vestergren Microm. Rar. Sel. 1703 (II).
France: Cote d’Or, Nov. 1889, F. Fautrey, Roum. Fg. Sel. Gall.
5212 (II). — April 1895, F. Fautrey, Roum. Fg. Sel. Gall.
7084 (II). — Hohneck, Alsace, July 1910, H. Sydow, Syd.
Ured. 2399 (II). — Cherbourg, Sept. 1924, G. Malenqon,
Herb. Bur. PI. Ind., U. S. Dept, of Agr., Washington, D. C.
(II). — Axat, Aude, April 30, 1932, G. D. Darker and G.
Fenwick-Owen (ii, in).
Great Britain: M. C. Cooke Fg. Brit. 2nd ed. (issued 1875) 73
(II). — Newton Abbot, England, April 1925, E. M. Wake¬
field, Kew Herb. (II, III). — Forden, England, May 1874,
John E. Vise, Herb. Vise Fg. Brit. 143 (II).
Italy: Gardone, May 25, 1912, O. Jaap, Jaap Fg. Sel. 571 (II).
Poland: Ojcow, undated, issued June 1911, B. Namyslowski, Herb.
Raciborski Myc. Polon. 186 (II). — Ojcow, July 25, 1925,
Wanda Konopacka, Herb. W. Siemaszko (II).
Caucasus: Kelassuri, near Suchum, May 10, 1915, W. Siemaszko,
Herb. Bur. PI. Ind., U. S. Dept, of Agr., Washington, D. C.
(II).
Milesia Scolopendrii is widely distributed throughout Eu¬
rope on its one known host Scolopendrium vulgare, and it ap¬
pears to be the first Milesia to receive a name. Fuckel (20)
described the uredo stage of this rust in 1873 as a Pyrenomy-
cete under the name Ascospora Scolopendrii, n. sp. To him the
uredinia were perithecia and the spores were asci, remarkable
because of their spiny surface. He interpreted granules in the
cytoplasm of his “asci” as spores too small to be measured.
He observed the peridium and commented on the white ex¬
trusions (“asci”). Oudemans (39) called attention to the
errors in Fuckel’s interpretations, both with respect to spores
and peridium. He concluded that the fungus in question was
a rust, and that it should be called uAecidium Scolopendrii,
n. comb.” or uEndophyllum Scolopendrii, n. comb.” It was
116
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
long cited as occurring on various other fern hosts. More
careful observations, however, have shown that the rusts on
these ferns were not M. Scolopendrii, and so one by one they
were described and re-classified.
In passing I should perhaps call attention to Vestergren’s
confused statements with reference to nos. 1702 and 1703 of
his Micromycetes rariores seleeti. The former is Milesia
Polypodii White \Milesina Dieteliana (Syd.) Magn.] on
Polypodium vulgare, but most of the synonyms cited are those
of M. Scolopendrii. The latter is Milesia Scolopendrii
(Fuckel) Arthur on Scolopendrium vulgare , which he desig¬
nates as Milesina Dieteliana — an untenable interpretation;
with historical inaccuracy he adds “N.B. Gehort wahrschein-
lich zu derselben Art wie Uredo Scolopendrii (Fuck.) Schroet.
auf Polypodium vulgare.”
So far there has been no suggestion that M. Scolopendrii
may comprise more than one recognizable, morphological
strain; but my examination of available material leads me to
suggest that such is not beyond the range of reasonable possi¬
bility. Thus a comparison of a collection from the Cote d’Or
in France with one from the Vosges shows that in one the
uredospores average 18 x 42 p with a wall less than one micron
thick, while those from the other average 18 x 35 p with a
wall twice the thickness of the former. Whether or not such
differences are constant could, of course, be determined only
by sustained observation under regularized culture methods.
A comparison of uredospores examined by me from various
collections is presented in Table 9. The averages given are
of 40 measured spores each, taken as they came in the mounts.
TABLE 9
Average
size of
Range in size
Collection
Uredospores
of Uredospores
Roum. Fung. Gall. 5212 (France)
18 x 42 p
14-20 x 28-57 p
Syd. Ured. 2399 (France)
18 x 35
16-23 x 28-45
Jaap Fung. Sel. 571 (Italy)
20 x 37
17-23 x 31-45
Herb. Siemaszko (Caucasus)
19 x 30
18-22 x 25-36
Racib. Myc. Polon. 186 (Poland)
18 x 39
15-22 x 30-49
Vise Fg. Brit. 143 (England)
21 x 46
18-25 x 36-58
Collection by Malengon (France)
20 x 32
16-23 x 25-44
Type (Switzerland)
19 x 33
16-22 x 25-41
Average of 320 spores
19 x 37 p
1032]
FAULL, THE GENUS MILESIA
117
Besides its European distribution, M. Scolopendrii is also
reported from Japan. Through the courtesy of Drs. Kamei
and Hiratsuka, I have had the opportunity of examining three
collections from Japan. In all of these the uredospores are
distinctly less strongly spinulose than in any of the European
gatherings I have seen; indeed, the spores in them are often
almost smooth. In view of this rather striking variation,
along with the occasional appearance of somewhat differently
shaped, more or less subapiculate spores in the Japanese mate¬
rial, I have thought it advisable to recognize the latter as a
variety of the species, namely, M. Scolopendrii var. sublevis ,
n. var. Finally M. Scolopendrii occurs in the Caucasus of Asia
according to collections made by Professor W. Siemaszko.
Although M. Scolopendrii has been repeatedly collected in
Europe during the last fifty years and more, there has been
no record heretofore of its teliospores. They are, however,
doubtless of frequent occurrence, but only on overwintered
fronds, in which development takes place at- about the time
the alternate host is in a stage susceptible to infection. Such
materials received from Dr. Eugene Mayor, Dr. G. D. Darker
and the Kew Herbarium, carry them in abundance, many of
the spore cells in some specimens with mature basidia, and it
is from these materials that my description of them has been
drawn. What the alternate host is remains to be demonstrated,
but in all likelihood it will be found to be Abies. With telial
material in hand little difficulty would be anticipated in estab¬
lishing the complete life cycle of this rust.
I am especially indebted to Professor Robert Chodat of the
University of Geneva for the opportunity of examining
Fuckers type material.
33. Milesia Scolopendrii (Fuckel) Arthur var. sublevis Faull,
var. nov. II et III.
Milesina Scolopendrii Hiratsuka in Jap. Jour. Bot. 3:313
(1917), in part, not Jaap. II and III.
Pycnidia et aecidia ignota. Sori uredosporiferi hypophylli,
epidermide tecti, sparsi vel aggregati, pustulati, rotundati,
0.1 -0.3 mm. diam., peridio ex cellulis hyalinis peridio inf erne
ex cellulis radiatim elongatis superne irregulariter polygonali-
bus composito cincti; cellulae peridii imbricatae, 7-17 p diam.,
118
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
pariete usque 1 m crasso. Uredosporae obovatae vel ellipsoi-
deae, rarius subapiculatae, subtile verruculosae, hyalinae,
17-23 x 28-54 p, plus minusve circa 19 x 38 p, episporio usque
1 p crasso. Teleutosporae intra cellulas epidermidis evolutae,
amphigenae, praecipue hypophyllae, verticaliter septatae,
pluricellulares, leves, hyalinae; cellulae teleutosporarum 8-17
x 9-25 p.
Hab. in foliis Scolopendrii vulgaris in Japonia.
O and I. Spermogonia and aecia unknown.
II. Uredinia as in the species; uredospores colorless, obo-
void or ellipsoid, occasionally subapiculate, 17-23 x 28-54 p,
averaging about 19 x 38 p; spore wall 0.5 to 1.0 p thick, with
fine, dot-like warts, rarely with a spine or two at the apex.
III. Telia on overwintered fronds, hypophyllous, occasion¬
ally epiphyllous, on indefinite, extensive pale yellow areas,
often involving entire fronds; teliospores within the epidermal
cells, one to a few in a cell, hyaline, one- to many-celled,
rounded or conforming to the shape of the containing cell,
anticlinally septate; the individual cells of the teliospores with
thin, smooth, colorless walls, irregularly polygonal except
along their free margins, 8-17 x 9-25 p.
Hosts and Distribution:
O and I. Unknown.
II and III. Scolopendrium vulgare Smith \Phyllitis Scolo-
pendrium (L.) Newm.l, in Japan. (See Hiratsuka in Jap.
Jour. Bot. 3:313. 1927.)
Type Locality: Mt. Teine, Ishikari, Japan, on Scolopendrium
vulgare. II and III.
Illustrations: Plate VI, Fig. 23.
Specimens Examined, all on Scolopendrium vulgare. —
Japan: Mt. Teine, Ishikari, June 17, 1925, Naohide Hiratsuka,
overwintered fronds (II, III); part of type. — Mt. Teine,
Ishikari, Oct. 22, 1924, Senji Kamei (II). — Maruyama, Sap¬
poro, Oct. 9, 1925, Naohide Hiratsuka (5732) (II).
Professor Naohide Hiratsuka (24) described for the first
time the teliospores of Milesia Scolopendrii var. sublevis. His
specimens were on overwintered fronds collected “in the spring
of 1925 at Maruyama and Mt. Teine near Sapporo,” Japan.
He stated that the telia cover “the whole surface of the leaves,”
and that the spores are intracellular. I find in material kindly
1932]
FAULL, THE GENUS MILESIA
119
shared by him that the teliospores are not aptly described as
being “divided into 2 to 4 cells.” Just as in other species they
are commonly divided into an indefinite and often large num¬
ber of cells, at times a single multicellular spore completely
filling an epidermal cell, conforming in outline to the latter.
FERN HOSTS AND THE SPECIES OF MILESIA THAT OCCUR
ON THEM
Fern Hosts
Species of Milesia
Magnusiana
Asplenii-incisi
magellanica
murariae
Feurichii
australis forma typica
australis forma irregularis
(O, I)
Blechni
Darken
Dennstaedtiae
andina
Miyabe
carpatica var. erythrosora
carpatica
Asplenium Adiantum nigrum L . M.
Asplenium incisum Thunb . M.
Asplenium magellanicum Kaulf . M.
Asplenium Ruta muraria L . M.
Asplenium septentrionale (L.) Hoffm. ..Af.
Blechnum auriculatum Cav . M.
Blechnum occidentale L . M.
Blechnum Spicant (L.) With . M.
Cryptogramma acrostichoides R. Br. . .M.
Dennstaedtia rubiginosa (Kaulf.)
Moore . M.
Dryopteris spec . M.
Dryopteris crassirhizoma Nakai . M.
Dryopteris erythrosora (Eaton)
Kuntze . M.
Dryopteris Filix mas (L.) Schott . M.
Dryopteris Filix mas (L.) Schott . M. Kriegeriana
Dryopteris marginalis (L.) A. Gray . . M. marginalis
Dryopteris patens (Sw.) Kunze . M . consimilis
Dryopteris spinulosa (O. F. Muller)
Kuntze . M. Kriegeriana
Dryopteris spinulosa var. americana
(Fisch.) Fernald . M.fructuosa
Dryopteris spinulosa var. dilatata
(Hoffm.) Underw . M. dilatata
Dryopteris spinulosa var. dilatata
(Hoffm.) Underw . M . Kriegeriana
Dryopteris spinulosa var. fructuosa
(Gilbert) Trudell . M.fructuosa
Dryopteris spinulosa var. intermedia
(Muhl.) Underw . M. intermedia
Dryopteris viridescens (Bak.) Kuntze . .M. Dryopteridis
Histiopteris incisa (Thunb.) J. Sm. . . .M. Histiopteridis
Nephrolepis cor difolia (L.) Presl . M. tenuis
Nephrolepis pendula (Raddi) J. Sm. . ,M . columbiensis
Nephrolepis rivularis (Vahl) Mett. . . ,M . insularis
Pellaea hastata (Thunb.) Prantl . M. nervisequa
Poly podium calif ornicum Kaulf . M . laeviuscula
forma typica
Polypodium glycyrrhiza D. C. Eaton . .M. laeviuscula
forma glycyrrhiza
Polypodium lycopodioides L . M. nervisequa
Poly podium virginianum L .
Polypodium vulgare L .
Polypodium vulgare L .
Polystichum aculeatum (L.) Roth
O, I,
(O, I)
Known
Stages
II
II
II
II, III
II, III
II
II, III
II, III
II, III
II
II
II, III
II, III
II, III
II, III
II, III
II
(I) II, III
II, III
II, III
II, III
(O, I)
(O)
,M. polypodophila
,M. Polypodii
. M . jezoensis
.M. Whitei
(O, I)
(O)
II, III
II, III
II, III
II
II
II
II
II
II, III
II, III
II
II, III
II, III
II, III
II, III
120
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Polystichum aculeatum (L.) Roth _ M. vogesiaca forma typica II, III
Polystichum aculeatum var. retrorso-
paleaceum Kodama . . . M . exigua II, III
Polystichum Braunii (Spenner) Fee ...M. exigua (O, I) II, III
Polystichum japonicum (Franch. et
Sav.) Diels . M. exigua II, III
Polystichum lob atum (Hudson) Presl . .M. vogesiaca forma typica II, III
Polystichum Lonchitis (L.) Roth . M. vogesiaca forma typica II
Polystichum munitum (Kaulf.) Presl . .M . Polystichi II, III
Polystichum munitum (Kaulf.) Presl. .M. vogesiaca forma typica II, III
Polystichum tripteron (Kunze) Pr. . . .M. exigua II, III
Polystichum varium (L.) Presl . M. vogesiaca
forma nipponensis II, III
Scolopendrium vulgare Smith (1793) M. Scolopendrii II, III
Scolopendrium vulgare Smith (1793) ..M. Scolopendrii var. sublevis II, III
THE SPECIES OF MILESIA, THE TWO GENERATIONS OF WHICH
ARE KNOWN, AND THEIR ALTERNATE HOSTS
Milesia Blechni
Milesia Kriegeriana
Milesia intermedia
Milesia marginalis
Milesia polypodophila
Milesia jezoensis
Milesia Miyabei
Milesia Dryopteridis
Milesia exigua
O, I Host
A bies cephalonica
Abies alba
Abies alba
Abies balsamea
Abies balsamea
Abies balsamea
A bies Mayriana
Abies Mayriana
Abies Mayriana
Abies Mayriana
A bies firma
Abies sachalinensis
II, III Host
Blechnum Spicant
Dryopteris spinulosa
Dryopteris spinulosa
var. intermedia
Dryopteris marginalis
Poly podium virginianum
Polypodium vulgare
Dryopteris crassirhizoma
Dryopteris viridescens
Polystichum Braunii
SPECIES OF MILESIA AND THEIR FERN HOSTS
(^Stages marked with an asterisk are reported or described for first time)
Milesia andina Faull . Dryopteris spec. II*
M. Asplenii-incisi Faull . Asplenium incisum II*
M. australis Arthur
forma typica Faull . Blechnum auriculatum II
forma irregularis Faull . Blechnum occidentale II, III
M. Blechni (Syd.) Arthur . Blechnum Spicant (O, I) II, III
M. carpatica (Wrob.) Faull . Dryopteris Filix mas II, III*
M. carpatica var. erythrosora Faull . . .Dryopteris erythrosora II*, III*
M. columbiensis (Diet.) Arthur . Nephrolepis pendula II
M. consimilis Arthur . Dryopteris patens II
M. Darken Faull . Crypto gramma acrostichoides II*, III*
M. Dennstaedtiae (Diet.) Faull . Dennstaedtea rubiginosa II
M . dilatata Faull . Dryopteris spinulosa
var. dilatata II*, III*
M. Dryopteridis (Kamei) Faull . Dryopteris viridescens (O) II, III
M. exigua Faull . Polystichum Braunii (O, I) II, III
Polystichum tripteron II*, III*
Polystichum aculeatum
var. retrorso-paleaceum II*, III*
Polystichum japonicum II*, III*
1932] FAULL, THE GENUS MILESIA
121
M . Feurichii (Magnus) Faull . Asplenium septentrionale II, III
M. fructuosa Faull . Dryopteris spinulosa
var. fructuosa II*, III*
Dryopteris spinulosa
var. americana II*, III*
M. Histiopteridis (Cunninghami) Faull .Histiopteris incisa II
M . insularis Faull . Nephrolepis rivularis II*
M. intermedia Faull . Dryopteris spinulosa
var. intermedia (0, I) II*, III*
M. jezoensis (Kamei et Hirat.) Faull . .Polypodium vulgare (0) II, III
M . Kriegeriana (Magnus) Arthur . Dryopteris Filix mas II, III
Dryopteris spinulosa (I) II, III
Dryopteris spinulosa
var. dilatata II, III*
M. laeviuscula (D. and H.) Faull
forma typica Faull . Polypodium calif ornicum II, III
forma glycyrrhiza Faull . Polypodium glycyrrhiza II, III*
M. magellanica Faull . Asplenium magellanicum II*
M. Magnusiana (Jaap) Faull . Asplenium A diantum nigrum II
M. marginalis Faull and Watson . Dryopteris mar ginalis (O, I*) II*, III*
M. Miyabei (Karoei) Faull . . Dryopteris crassirhizoma (O, I) II, III
M. murariae (Magnus) Faull . Asplenium Ruta muraria II, III*
M. nervisequa (von Thiimen) Faull . . .Pellaea hastata II
Poly podium lycopodioides II
M. Polypodii B. White . Poly podium vtdgare II, III
M. polypodophila (Bell) Faull . Polypodium virgimanmn (O, I) II, III
M. Polystichi Wineland . Polystichum munitum II, III*
M. Scolopendrii (Fuckel) Arthur . Scolopendrium vulgare II, III*
M. Scolopendrii var. sublevis Faull _ Scolopendrium vulgare II*, III
M. tenuis Faull . Nephrolepis cor difolia II*
M. vogesiaca (Sydow) Faull
forma typica Faull . Polystichum aculeatum II, III*
forma typica Faull . Polystichum lobatum II, III*
forma typica Faull . Polystichum Lonchitis II*
forma typica Faull . Polystichum munitum II*, III*
forma nipponensis Faull . Polystichum varium II*, III*
M. Whitei Faull . Polystichum aculeatum II*, III*
EXCLUDED SPECIES
1. Milesina Lygodii (Har.) Sydow.
The first record of a rust on Lygodium is to be found in a
paper by Hariot (Jour. Bot. 14:117. 1900); this rust he
called Uredo Lygodii, n. sp. The species is based on uredinial
material collected on Lygodium spec, at Pernambouco, Brazil.
(Type in Herb. Mus. Paris, no. 1229, undated, collector M.
Gardener).
The next collection of a rust on Lygodium of which I find
any account was made by J. N. Rose and P. G. Russell near
Bahia, Brazil, May 28, 1915, on Lygodium polymorphum
(Cav.) H.B.K. Two packets under number 21514 are pre¬
served, one in the Herbarium of the Carnegie Institution of
Washington, and the other in the Herbarium of the New York
Botanical Garden; a third packet, under number 19664 a, is
122
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
in the U. S. National Herbarium, Smithsonian Institution.
This collection was studied by Arthur (4), who found two
types of spores which he designated uredospores and telio-
spores respectively; he referred the species to Puccinia under
the name Puccinia ( Dicaeoma ) Lygodii (Har.) Arthur, n.
comb. Arthur considered that the earlier collection by Har-
iot was the same species as the one made by Rose and Russell.
A third collection (uredinial) is preserved in the Farlow
Herbarium of Harvard University under the name Uredo
Lygodii Har., determined by Arthur. It was made by Profes¬
sor Roland Thaxter at St. Anna Valley, Trinidad, Feb. 20,
1913, on Lygodium polymorphum (Cav.) H.B.K.
A fourth collection (uredinial) I found in the Arthur Herbar¬
ium, Purdue University, under the name Milesia Lygodii. This
was made by E. W. D. and Mary M. Hoi way at Rio de Janeiro,
Brazil, Aug. 27, 1921, on Lygodium volubile Sw.
A fifth collection (uredinial) is to be found in Paul C.
Standley’s collection (19193), “Plants of El Salvador/’ made
in the vicinity of San Salvador, Dec. 20-Jan. 4, 1922, on
Lygodium polymorphum (Cav.) H.B.K. This was determined
by Arthur (3) as Milesia Blechni (Syd.) Arthur, n. comb.,
but listed by him in the North American Flora (5) under the
name Milesia australis Arth.
Last of all, a collection (uredinial) on Lygodium spec, made
by Professor F. L. Stevens (no. 154) at Tumatumaris, British
Guiana, July 11, 1922, was studied by Sydow who named it
Milesina Lygodii Syd., n. sp. (H. Sydow, Rusts of British
Guiana and Trinidad. Mycologia, 17:255. 1925).
I have had the opportunity of making a comparative ex¬
amination of specimens from all of these collections. I find
that their uredinia are much alike with regard to size, aspect,
habit, peridia and spore characters. Unfortunately there is no
record of spore color for any one of them. The existence of
stalked, septate spores in the second collection referred to
above, interpreted by Professor Arthur as “teliospores,” would
appear to make its inclusion in the genus Milesia untenable.
But if we exclude that collection the others also should be ex¬
cluded because of general uredinial similarity throughout.
Further studies of these rusts on Lygodium is needed before a
final decision can be reached as to just where they belong.
1932]
FAULL, THE GENUS MILESIA
123
2. Milesina Pteridis (D. & H.) Sydow in Monog. Ured.
3:481 (1915).
Dietel (10) described Uredo Pteridis D. and H. in 1894
from material collected by A. J. McClatchie at Pasadena,
Calif., January 1893, on uPteris aquilina.” Sydow (44)
transferred the species to Milesina and firmly maintained that
it had nothing to do with Uredinopsis Pteridis D. and H. He
cited specimens 2988 and 4087 in Bartholomew’s Fungi Co-
lumbiani, and number 881 in Bartholomew’s North American
Uredinales as examples of Milesina Pteridis. On examina¬
tion of the last two (parts of an identical collection from
British Columbia) I find typical subepidermal teliospores of
a Uredinopsis , and this is the only telial-bearing material
Sydow mentions. The specimens of the first named collection
carry uredinia only, but their uredospores are similar to those
in the others cited. Therefore, Sydow’s contention of the
existence of a Milesina Pteridis is not supported by any evi¬
dence so far adduced, and in part at least is positively dis¬
proved.
3. Milesia mirabilis (Pk.) Arthur (1) = Uredinopsis ameri-
cana Syd.
4. Milesia Atkinsonii (Magn.) Arthur (1 )— Uredinopsis
Atkinsonii Magn.
5. Milesia Pteridis (D. and H.) Arthur {Y) — Uredinopsis
Pteridis D. and H.
6. Milesia Osmundae (Magn.) Arthur (1 ) — Uredinopsis
Osmundae Magn.
7. Milesia Copelandi (Syd.) Arthur (1) = Uredinopsis Cope-
landi Syd.
ACKNOWLEDGMENTS
It is with pleasure that I acknowledge helpful contacts with
Professor J. C. Arthur, Dr. C. L. Shear, Professor Eugene
Mayor, Professor Naohide Hiratsuka, Mr. Alfred Rehder, Mr.
George B. Cummins, Professor H. S. Jackson and Professor
W. C. Blasdale during the course of my study of the genus
Milesia; opportunity to examine the collections of the follow¬
ing Herbaria through the courtesy of their respective Cura¬
tors, — Farlow, Arthur (Purdue University), N. Y. Agricul¬
tural College, N. Y. State Museum (Albany), University of
124
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Wisconsin, Oregon State College, University of Washington,
University of California, U. S. Bureau of Plant Industry,
N. Y. Botanical Garden, Connecticut Agricultural College,
Kew, British Museum, Musee d’Histoire Naturelle (Paris),
Universite de Geneve and the Arnold Arboretum; contributions
of specimens from most of the foregoing, also from Professor
Roland Thaxter, Director A. Wroblewski, Mr. W. R. Watson,
Professor Lee Bonar, Mr. H. E. Parks, Professor H. P. Bell,
Professor H. W. Thurston, Jr., Dr. W. Wollenweber, Mr.
J. R. Hansbrough, Mr. T. W. Childs, Professor W. Siemaszko,
Dr. E. M. Doidge and Dr. S. Kamei; data regarding the O-
stage from Miss L. M. Hunter; advice from Mr. C. A. Weath-
erby with respect to the nomenclature of the fern hosts; aid in
collecting and constant interest on the part of my Assistant,
Dr. G. D. Darker.
SUMMARY
1. The genus Milesia was founded by Dr. F. B. White
in 1877 on the uredinial stage of a new species, M. Polypodii
White, a rust occurring on Poly podium vulgar e L. in Europe.
The same species was described by Sydow in 1903 under the
name Melampsorella Dieteliana, n. sp. Magnus without suf¬
ficient reason changed the name of the genus to Milesina in
1909.
2. The species of Milesia are quite readily separated from
one another by their remarkably versatile uredospores and
other uredinial characters. Their teliospores are almost non-
distinctive specifically.
3. Consonant with the accepted definition of the genus,
Milesia is defined in this paper as a genus of white-spored
rusts, characterized by immersed, hemispherical spermogonia,
peridermia, uredinia with a peridium and non-catenulate
uredospores, and many-celled, intraepidermal teliospores.
4. Species of Abies are the aecial hosts of the nine species
of Milesia for which the aecial hosts have been determined.
5. Peridermium balsameum Peck in America and Aecidium
pseudo columnar e Kuehn in Europe, as generally understood,
have included species of Milesia as well as of Uredinopsis.
6. Species of the Polypodiaceae are the sole hosts of the
1932]
FAULL, THE GENUS MILESIA
125
uredinial and telial stages of the species of Milesia. Most of
the subfamilies of the Polypodiaceae are included in this con¬
nection.
7. Species of Milesia are continued from one growing sea¬
son to the next, that is, “overwintered,” on the fern hosts. A
copious crop of uredospores is produced on the overwintered
fronds. Apparently a species may be continued indefinitely
from year to year without reference to an aecial host. Con¬
ceivably this may mean the possibility of some evolution of
new species within the genus without involving the aecial
phase.
8. The teliospores of a few species are regularly produced
on fronds of the current season, but in most species so far as is
known their formation is on overwintered fronds only in the
spring. They occur in great abundance and not rarely or
scantily as has been supposed.
9. The aecia, where known, are annual on Abies and form
on needles of the current season only — with one exception,
Milesia polypodophila. M. polypodophila is perennial in the
stems and needles of Abies, causes loose brooms and its aecia
are to be found on needles 3-9 years of age.
10. The geographical range of Milesia is world-wide. Nine
species have been collected in the United States and Canada,
seven in Central America, the West Indies and tropical South
America, eleven in Europe, two in Africa (one in the north
and one in the south), thirteen in Asia (four in the west and
nine in the east), and one in Australasia. One species only
found beyond the range of Abies is known to produce telio¬
spores, namely, M. australis in Colombia.
11. One species only is known to occur in both the Eastern
and the Western Hemispheres, namely, , Milesia vogesiaca
(Syd.) Faull. The fern hosts in this case are species of
Poly stic hum; but the hosts in the two Hemispheres respec¬
tively are different species of Polystichum. There is a possi¬
bility that M. vogesiaca is a palingenetic species.
12. Thirty-three species and two varieties of Milesia are
recognized in this paper.
13. Thirteen species and varieties are described as new,
126
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
namely, M. andina, M. Asplenii-incisi, M. carpatica var.
erythrosora, M. Darken , M. dilatata, M. fructuosa, M. insu-
laris, M. intermedia, M. magellanica, M. marginalis, M.
Scolopendrii var. sublevis, M. tenuis and M. Whitei. One is
transferred from the genus Hyalopsora, namely, Milesia
laeviuscula (D. and H.) Faull.
14. The telial stage of twenty-four species and varieties of
Milesia are described in this paper. The teliospores of the
following are described for the first time — M. Darken, M.
dilatata, M. carpatica, M. carpatica var. erythrosora, M. fruc¬
tuosa, M. intermedia, M. marginalis, M. Polystichi, M. Whitei,
M. Scolopendrii, M. murariae and M. vogesiaca (without a
qualifying question mark).
15. Species of Milesia are apparently not closely restricted
with respect to their aecial hosts. They are in general closely
restricted with respect to their uredinial and telial hosts.
Laboratory of Plant Pathology, Arnold Arboretum,
Harvard University.
BIBLIOGRAPHY
1 . Arthur, J. C. Eine auf die Struktur und Entwickelungsgeschichte
begriindete Klassification der Uredineen. [Resultats
Scientifiques du Congres Internationale de Botanique.
Vienna (1905). 1906.]
2. - Uredinales of Porto Rico based on collections by F.
L. Stevens. [Mycologia, 7:168-196 (continued). 1915.]
3. - Uredinales collected by R. Thaxter and J. B. Rorer in
Trinidad. (Bot. Gaz. 73:58-69. 1922.)
4. - New species of Uredineae (XV). (Bull Torr. Bot.
Club 51:51-59. 1924.)
5. - Uredinales (of North America) — Milesia. (North
American Flora, 7:685-68 7. 1925.)
6. Bell, H. P. Fern rusts of Abies. (Bot. Gaz. 77:1-31. 1924.)
7. Blasdale, W alter C. A preliminary list of the Uredinales of
California. (Univ. of Calif. Publications in Botany,
7:101-157. 1919.)
8. Bubak, F. Houby Ceske. Dil 1. Rezy (Uredinales) : 207-8. 1906.
9. Cunningham, G. H. The Uredinales, or rust fungi, of New
Zealand. Supplement to Parts 1 and 2. (Trans. N. Z.
Inst. 55:1-58. 1924.)
10. Dietel, P. New California Uredineae. II. (Erythea, 2:127. 1894.)
11. - Ueber die Teleutosporenform von Uredo laeviuscula
D. et H. und iiber Melampsora Fagi D. et Neg. (Annal.
Mycol. 1:415-417. 1903.)
1932]
FAULL, THE GENUS MILESIA
127
12. - in Eug. Mayor’s “Contribution a l’etude des Uredinees
de Colombie.” (Mem. Soc. Sci. Nat. Neuchatel 5:422-
599. 1914.)
13. Dietel, P. & F. Neger. Uredinaceae chilensis 1. (Bot. Jahrb.
22:348-358. 1897.)
14. Doidge, Ethel M. A preliminary study of the South African
rust fungi. (Bothalia, 2:1-228. 1927.)
15. Faull, J. H. Balsam rusts. (Rept. Minister of Lands and For¬
ests of the Province of Ontario for 1922: 253-4. 1923.)
16. - The morphology, biology, and phylogeny of the Puc-
ciniastreae. [Proc. Internat. Congr. Plant Sciences
(Ithaca, 1926) 2:1735-1745. 1929.]
17. - Notes on forest diseases in Nova Scotia. (Jour.
Arnold Arb. 9:55-58. 1930.)
18. - Milesina rusts on Aspidium Braunii Spenner. (Jour.
Arnold Arb. 12:218-9. 1931.)
19. Fischer, Ed. Die Uredineen der Schweiz. Bern. 1904.
20. Fuckel, L. Symbolae Mycologicae. Beitrage zur Kenntniss der
Rheinischen Pilze. Zweiten Nachtrag. Wiesbaden. 1873.
21. Grove, W. B. The British rust fungi. Cambridge. 1913.
22. - The British species of Milesina. (Jour. Bot. 59:109.
1921.)
23. - Mycological notes VI. (Jour. Bot. 59:311-313. 1921.)
24. Hiratsuka, N. A contribution to the knowledge of the Melamp-
soraceae of Hokkaido. (Jap. Jour. Bot. 3:289-322.
1927.)
25. Hotson, J. W. Preliminary list of the Uredinales of Washing¬
ton. (Publications of the Puget Sound Biological Sta¬
tion of the Univ. of Wash. 4:273-391. 1925.)
26. Hunt, W. R. The Uredinales or rusts of Connecticut. (State
Geol. and Nat. His. Survey, Bull. 36:178. 1926.)
27. Hunter, L. M. Comparative study of spermogonia of rusts of
Abies. (Bot. Gaz. 83:1-23. 1927.)
27a. - The identity of Peridermium balsameum Peck. (Manu¬
script.)
28. Jackson, H. S. The Uredinales of Oregon. (Brookl. Bot. Gard.
Mem. 1:198-297. 1918.)
29. Kamei, Senji. Notes on Milesina vogesiaca Sydow on Polysti-
chum Braunii Fee and its peridermial stage on the
needles of Abies Mayriana Miyabe et Kudo, A. firma
Sieb. et Zucc. and A. sachalinensis Mast. (Trans. Sap¬
poro Nat. His. Soc. 11:141-147. 1930.)
30. - A new species of Milesina parasitic on Polypodium
vulgare L. (Trans. Sapporo Nat. His. Soc. 12:27-33.
1931.)
31. Klebahn, H. Kryptogamenflora der Mark Brandenburg 5a,
5:851-2. 1914.
Kulturversuche mit Rostpilzen. (Zeitsch. Pflanzenkr.
26:257-277. 1916.)
32.
128
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
33. Liro (Lindroth), J. Ivar. Uredineae Fennicae. Helsingfors.
1908.
34. Magnus, P. Die Teleutosporen der Uredo Aspidiotus Peck. (Ber.
Deutsch. Bot. Ges. 13:285-288. 1895.)
35. - Weitere Mitteilungen fiber die auf Farnkrautern auf-
tretenden Uredineen. (Ber. Deutsch. Bot. Ges. 19:578-
584. 1901.)
36. - M damps or ellci Feurichii, eine neue Uredinee auf
Asplenium septentrionale. (Ber. Deutsch. Bot. Ges. 20:
609-612. 1902.)
37. - - Bemerkungen fiber einige Gattungen der Melampso-
reen. (Ber. Deutsch. Bot. Ges. 27:325. 1909.)
38. Moss, E. H. The uredo stage of the Pucciniastreae. (Ann. Bot.
40:813-847. 1926.)
39. Oudemans, C. A. J. A. Drei unrichtig bestimmte Pilze. (Bot.
Zeitung, 33:585-592. 1875.)
40. Rostrup, E. Mykologiske Meddelelser (VII). Spredte Iagttagelser
fra 1895-1896. (Bot. Tidsskr. 21:37-49. 1897-98.)
41 . Schroeter, J. in Cohn, Kryptogamen-Flora von Schlesien. Band
3, Erste Halfte, p. 374. 1889.
42. Sydow, H. & P. Mycotheca germanica, Fasc. 2 (nos. 51-100).
(Annal. Mycol. 1:536-539. 1903.)
43. - Mycotheca germanica, Fasc. 18-19 (nos. 851-950).
(Annal. Mycol. 8:489-493. 1910.)
44. - Monographia Uredinearum, 3:473-481. 1915.
45. - Mykologische Mitteilungen. (Annal. Mycol. 16:240-
248. 1918.)
46. Von Thumen, F. Fungi Austro-Africani (V). (Flora, 60:407-
413. 1877.)
47. Thurston, Jr., H. W. An interesting fern rust new to the
United States. (Mycologia, 20:44-45. 1928.)
48. White, F. Buchanan. Note on the zoology and botany of Glen
Tilt. (Scottish Naturalist, 4:160-163. 1877-8.)
49. - On Uredo filicum Desm. and Uredo (?) pteridium
n. sp. (Scottish Naturalist 4:27. 1877-8.)
50. Wroblewski, A. Przyczynek do znajomosci grzybow Pokucia.
(Sprawozdania komisyi fizyograficzna, Akademija
umiejetnosci, Krakow, 47:147-178. 1913.)
51. Hiratsuka, N. & Y. Yoshida. Two species of Milesina on some
Japanese species of Polystichum. (Trans. Tottori Soc.
Agr. Sc. 4:7-10. 1932.)
52. Kamei. S. On new species of heteroecious fern rusts. (Trans.
Sapporo Nat. His. Soc. 12, 3:161-174. 1932.)
1932]
FAULL, THE GENUS MILESIA
129
ADDENDUM TO “V. SPECIES ON DRYOPTERIS”.
Just as the page proof of this paper was being corrected
there came to hand a reprint from Dr. Kamei (52) in which
are described two new species of Milesia. I have not seen
specimens of either, but for the sake of completeness I include
a synopsis of his descriptions under numbers 34 and 35. The
stages on Abies were described by Kamei from cultures, but
almost no data are given. The only cultures mentioned by
him for both species were from the fern hosts to Abies.
34. Milesia Miyabei (Kamei) Faull, n. comb. O, I, II and
III.
Milesina Miyabei Kamei in Trans. Sapporo Nat. His. Soc.
12, 3:169 (1932). O, I, II and III.
O. Spermogonia hypophyllous, few, inconspicuous, almost
spherical, 160-280 p broad by 190-240 p deep; spermatia hya¬
line, narrowly cylindrical 1. 6-2.0 x 4. 8-7. 6 p.
I. Aecia hypophyllous on needles of current season, white,
cylindrical, up to 2 mm. high by 0.5 mm. in diameter; peridium
colorless; peridial cells polygonal, overlapping, 17-26 x 23-41 p
with outer walls thin, smooth, with inner walls verrucose, 2-4 p
thick; * aeciospores globose or ellipsoid, colorless, verrucose,
14-24 x 15-29 p, averaging about 18 x 22 p.
II. Uredinia hypophyllous, subepidermal, scattered on
greenish to brown areas of indefinite extent, pustular, covered
by yellowish to brown epidermis, round, 0.15-0.35 mm. in
diameter; peridium subhemispherical, delicate; uredospores
colorless, extruded in delicate tendrils through a centrally
placed apical pore, fusiform, narrow, generally truncate at
apex, narrowed toward base, 11-20 x 26-55 p; walls of spores
about 1 p thick, smooth.
III. Telia amphigenous on overwintered fronds; telio-
spores within the epidermal cells, hyaline, smooth, few- to
many-celled.
Hosts and Distribution:
O and I. Abies Mayriana Miyabe and Kudo, in Japan.
(Cultures).
II and III. Dryopteris crassirhizoma Nakai, in Japan.
Kamei (51) informs us that the spermogonia of Milesia
Miyabei are very similar to those of M. polypodophila, though
130
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
“slightly wider.’7 He also points out that the uredospores are
“more or less bone-shaped” and very characteristic. The
teliospores are typically produced on overwintered fronds in
the early part of June.
35. Milesia Dryopteridis (Kamei) Faull, n. comb. O, II
and III.
Milesina Dryopteridis Kamei in Trans. Sapporo Nat. His.
Soc. 12, 3:171 (1932). O, II and III.
O. Spermogonia on needles of current season, hypophyl-
lous, minute, inconspicuous.
I. Unknown.
II. Uredinia hypophyllous, subepidermal, scattered on
rather restricted brownish discolored areas, pustular, round,
0.1 -0.25 mm. in diameter; peridium hemispherical, delicate;
peridial cells isodiametrically to irregularly polygonal, 8 x
15 p; uredospores colorless, extruded in delicate tendrils
through a centrally placed apical pore, obovoid or ellipsoid,
12 - 15 x 17-28 p, averaging about 13 x 22 p; walls of spores
distantly and delicately echinulate, 1.5- 2.5 p thick.
III. Telia on the fronds; teliospores within the epidermal
cells, 1- to many-celled, hyaline smooth.
Hosts and Distribution:
O. Abies Mayriana Miyabe and Kudo, in Japan. (Cul¬
tures) .
I. Not known, but certainly existent.
II and III. Dryopteris viridescens (Bak.) Kuntze, in
Japan.
The description of the uredinia and the uredospores of
Milesia Dryopteridis given by Kamei (52) at once calls to
mind M. carpatica. The main difference is in the thickness of
uredospore walls. So far as can be judged from Kamei’s meager
description of the teliospores their cells are larger than those
of M. carpatica. It is to be noted that M. carpatica var. ery-
throsora on Dryopteris erythrosora in Japan demands further
study before a final conclusion can be reached as to whether or
not it differs sufficiently from M. carpatica to warrant varietal
distinction. Likewise, judging from Kamei’s description of
M . Dryopteridis , I feel that it should be closely compared with
M. carpatica both as to morphology and host restrictions in
a closer check on its taxonomic status.
1932]
FAULL, THE GENUS MILESIA
131
EXPLANATION OF PLATES
Magnifications: Figs. 1-33, about 1050 times; Figs. 34-37, about 800
times.
PLATE I
Fig.
Fig.
Fig.
Fig.
1 a-d. Uredospores of Milesia Polypodii B. White. Syd. Ured.
2396.
2 a-d. Uredospores of M. laeviuscula (D. & H.) Faull, forma
typica. N. Am. Ured. 1505.
3 a-d. Uredospores of M. laeviuscula forma glycyrrhiza Faull.
Path. Herb. Arnold Arb. 333.
4 a-d. Uredospores of M. polypodophila (Bell) Faull. Herb.
J. H. Faull 6788.
Fig. 5 a-d.
Fig. 6 a-d.
Fig. 7 a-d.
Fig. 8 a-d.
Fig. 9 a-d.
Fig. 10 a-d.
Fig. 11 a-d.
Fig. 12 a-d.
Fig. 13 a-d.
Fig. 14 a-d.
Fig. 15 a-d.
Fig. 16 a-d.
PLATE II
Uredospores of M. Blechni (Sydow) Arthur, a, b, Syd.
Myc. Germ. 877 ; c, d, Syd. Ured. 2092.
Uredospores of M. australis Arthur forma typica. Syd.
Ured. 44a.
Uredospores of M. australis forma irregularis Faull.
a-c, Eug. Mayor, Colombia, Aug. 16, 1910; d, H. H.
Whetzel et al., Porto Rico, Jun. 22, 1924.
Uredospores of M. carpatica (Wrob.) Faull. A. Wrob-
lewski, Poland. From type.
PLATE III
Uredospores of M. Kriegeriana (Magnus) Arthur, a, d,
Syd. Ured. 1634; b, Krieg. Fg. Saxon. 856; c, Rabh.
Fg. Eur. 4242.
Uredospores of M. dilatata Faull. Path. Herb. Arnold
Arb. 284.
Uredospores of M. intermedia Faull. a-c, Herb. J. H.
Faull 9933; d, idem 7527.
Uredospores of M. marginalis Faull and Watson. Herb.
J. H. Faull 7526.
PLATE IV
Uredospores of M. Feurichii (Magnus) Faull. Krieg.
Fg. Saxon. 2008.
Uredospores of M. murariae (Magnus) Faull. Jaap Fg.
Sel. 530.
Uredospores of M. Magnusiana (Jaap) Faull. Jaap Fg.
Sel. 623.
Uredospores of M. magellanica Faull. R. Thaxter, Cor¬
ral, Chili, Dec. 1905.
132
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
PLATE V
Fig. 17 a-d. Uredospores of M. vogesiaca (Sydow) Faull. Path.
Herb. Arnold Arb. 171.
Fig. 18 a-d. Uredospores of M. Polystichi Wineland. Path. Herb.
Arnold Arb. 183.
Fig. 19 a-d. Uredospores of M. exigua Faull. A. Wroblewski, Po¬
land, Aug. 1913.
Fig. 20 a-d. Uredospores of M. Whitei Faull. E. M. Wakefield,
England, Apr. 1925. Kew. Herb.
PLATE VI
Fig. 21 a-d.
Fig. 22 a-c.
Fig. 23.
Fig. 24 a-d.
Uredospores of M. consimilis Arthur. L. M. Under¬
wood, Jamaica, April 22, 1903.
Uredospores of M. Scolopendrii (Fuckel) Arthur, a, b,
Syd. Ured. 2399; c, Roum. Fg. Sel. 5212.
Uredospore (occasional shape) of M. Scolopendrii var.
sublevis Faull. S. Kamei, Japan, Oct. 22, 1924.
Uredospores of M. Darken Faull. Path. Herb. Arnold
Arb. 254.
Fig. 25 a-d. Uredospores of M. Dennstaedtiae (Dietel) Faull. Eug.
Mayor, Colombia, Aug. 20, 1910.
PLATE VII
Fig. 26 a-d.
Fig. 27 a-d.
Fig. 28 a-d.
Fig. 29 a-d.
Uredospores of M. columbiensis (Dietel) Arthur, in
part. Eug. Mayor, Colombia. From type.
Uredospores of M. insularis Faull. F. L. Stevens, Porto
Rico, Nov. 25, 1913.
Uredospores of M. nervisequa (von Thuemen) Faull.
Thuem. Myc. Univ. 1141.
Uredospores of M. Asplenii-incisi Faull. Naohide
Hiratsuka, Japan, Mar. 26, 1930.
PLATE VIII
Fig. 30 a-d. Uredospores of M. andina Faull. G. Lagerheim, Ecua¬
dor, 1890.
Fig. 31 a-d. Uredospores of M. Histiopteridis (Cunningham) Faull.
Herb. G. H. Cunningham, New Zealand, 1318.
Fig. 32 a-d. Uredospores of M. tenuis Faull. Philippine Fungi,
Herb. Univ. Calif. 292430.
Fig. 33 a-d. Uredospores of M. jezoensis (Kamei) Faull. S. Kamei,
Japan, Sept. 30, 1930.
1932]
FAULL, THE GENUS MILESIA
133
PLATE IX
Fig1. 34. Teliospores of M. vogesiaca (Sydow) Faull. Eug.
Mayor, Switzerland, June 22, 1925.
Fig. 35. Teliospores of M. dilatata Faull. Path. Herb. Arnold
Arb. 284.
Fig. 36, 37. Teliospores of M. polypodophila (Bell) Faull. Herb.
J. H. Faull 4867.
INDEX
Synonyms are printed in italics and new names and new combinations in
bold-face type
ABIES
alba .
balsamea .
cephalonica
firma .
Mayriana .
sachalinensis
. 38, 59
. 66, 71, 90
. 38
. 101
88, 101, 129, 130
. 101
AECIDIUM
pseudocolumnare . 37
Scolopendrii . 113
ASCOSPORA
Scolopendrii . 113
ASPLENIUM
Adiantum nigrum . 33
incisum . 31
magellanicum . 32
Ruta muraria . 35
septentrionale . 27
BLECHNUM
auriculatum . 42
hastatum . 42
occidental . 42
Spicant . 38
CAEOMA
nervisequum . 77
CRYPTOGRAMMA
acrostichoides . 47
DENNSTAEDTIA
rubiginosa . 48
DRYOPTERIS
crassirhizoma . 129
erythrosora . 58
Filix mas . 56, 59
marginalis . 71
patens . 63
spinulosa . 59
spinulosa var. americana . 52
spinulosa var. dilatata . 50, 59
spinulosa var. fructuosa . 52
spinulosa var. intermedia . 66
spec. (aff. D. oligocarpa) . 55
viridescens . 150
HI STIOPTERIDIS
incisa .
74
136 CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
HYALOPSORA
Feurichii . 27
Kriegeriana . 59
laeviuscula . 95
LYGODIUM
polymorphum . . . 119
volubile . 119
MELAMPSORELLA
Blechni . 37
Dieteliana . 81
Feurichii . 27
Kriegeriana . 58
MILESIA . 22
andina (PI. VIII, fig. 30 a-d) . 25, 54
Asplenii-incisi (PI. VII, fig. 29 a-d) . 24, 30
Atkinsonii . 121
australis forma irregularis (PI. II, fig. 7 a-d) . 24, 41
australis forma typica (PI. II, fig. 6 a-d) . 24, 41
Blechni (PI. II, fig. 5 a-d) . 24, 37
carpatica (PI. II, fig. 8 a-d) . 25, 55
carpatica var. erythrosora . 25, 57
columbiensis (PI. VII, fig. 26 a-d) . 25, 75
consimilis (PI. VI, fig. 21 a-d) . 25, 63
Copelandi . 121
Darkeri (PI. VI, fig. 24 a-d) . 24, 46
Dennstaedtiae (PI. VI, fig. 25 a-d) . 24, 48
dilatata (PI. Ill, fig. 10 a-d; IX, fig. 35) . 25, 49
Dryopteridis . 130
exigua (PI. V, fig. 19 a-d) . 26, 100
Feurichii (PI. IV, fig. 13 a-d) . 24, 27
fructuosa . 25, 51
Histiopteridis (PI. VIII, fig. 31 a-d) . 25, 73
insularis (PI. VII, fig. 27 a-d) . 25, 76
intermedia (PI. Ill, fig. 11 a-d) . 25, 64
jezoensis (PI. VIII, fig. 33 a-d) . 26, 87
Kriegeriana (PI. Ill, fig. 9 a-d) . 25, 58, 64, 69
laeviuscula forma glycyrrhiza (PI. I, fig. 3 a-d; text fig1. 2) ... 26, 95
laeviuscula forma typica PI. I, fig. 2 a-d; text fig. 1) . 26, 95
Lygodii . 119
magellanica (PI. IV, fig. 16 a-d) . 24, 31
Magnusiana (PI. IV, fig. 15 a-d) . 24, 32
marginalis (PI. Ill, fig. 12 a-d) . 25, 69
mirabilis . 121
Miyabei . 129
murariae (PI. IV, fig. 14 a-d) . 24, 34
nervisequa (PI. VII, fig. 28 a-d) . 26, 77
Osmundae . 121
Polypodii (PI. I, fig. 1 a-d) . 26, 81
polypodophila (PI. I, fig. 4 a-d) . 26, 89
Polystichi (PI. V, fig. 18 a-d) . 26, 108
Pteridis . 121
pycnograndis . 89
Scolopendrii (PI. VI, fig. 22 a-c) . 26, 113
1932]
INDEX
137
Scolopendrii var. sublevis (PI. VI, fig. 23) . 26, 117
tenuis (PI. VIII, fig. 32 a-d) . 25, 74
vogesiaca (PI. V, fig. 17 a-d; IX, fig. 34) . 26, 103
vogesiaca forma nipponensisi . 108
Whitei (PI. V, fig. 20 a-d) . 26, 111
MILES1NA . 22
Blechni . 37, 41
carpatica . 55
columbiensis . 75
consimilis . 63
Dennstaedtiae . 48
Dieteliana . 81
Dryopteridis . 130
exigna . 100
Feurichii . 27
Histiopteridis . 73
jezoensis . 87
Kriegeriana . 59
Lygodii . 119
Magnusiana . 32
margincdis . 69
Miyabei . 129
Murariae . 34
nervisequa . 77
polypodophila . 89
Pteridis . 120
Scolopendrii . 113, 117
vogesiaca . 100, 103
NEPHROLEPIS
cordifolia . 75
pendula . 76
rivularis . 77
PELLAEA
hastata . 78
PERIDERMIUM
balsameum . 64
pycnogrande . 89
PHYLLITIS
Scolopendrium . 114, 118
POLYPODIUM
californicum . 96
glycyrrhiza . 96
lycopodioides . 78
virginianum . 91
vulgare . 82, 88
vulgare var. serratum . 82
POLYSTICHUM
aculeatum . 104, 112
aculeatum var. retrorso-paleaceum . 101
138
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM [No. 2
Braunii .
japonicum
lobatum
Lonchitis
munitum
tripteron
varium .
.... 101
. ... 101
. ... 104
. ... 104
104, 109
. ... 101
. ... 104
PUCCINIA
Lygodii . 119
pteridum . 113
SCOLOPENDRIUM
vulgare . 114, 118
THEKOPSORA
laeviuscula . . . 95
UREDINOPSIS
polypodophila . 89
Scolopendrii . 37, 113
UREDO
Blechni .
laeviuscula .
Lygodii .
murariae .
pteridium .
Scolopendrii . 34, 37,
vogesiaca .
41
95
119
34
113
113
103
Contrib. Arnold Arb. No. 2
Plate I
1.
3.
The Genus Milesia
M. Polypodii.
M. laeviuscula f. glycyrrhiza.
2. M. laeviuscula f. typica.
4. M. polypodophila.
Contrib. Arnold Arb. No. 2
Plate II.
The Genus Milesia
S. M. Blechni. 6. M. australis f. typica.
7. M. australis f. irregularis. 8. M. carpatica.
.
i
.
Contrib. Arnold Arb. No. 2
Plate III
The Genus Milesia
9. M. Kriegeriana. 10. M. dilatata.
11. M. intermedia. 12. M. marginalis.
Contrib. Arnold Arb. No. 2
Plate IV.
14
15
13. M. Feurichii.
IS. M. Magnusiana.
14. M. murariae.
16. M. magellanica.
Contrib. Arnold Arb. No. 2
Plate V
17. M. vogesiaca. 18. M. Polystichi.
19. M. exigua. 20. M. Whitei.
Contrib. Arnold Arb. No. 2 Plate VI.
The
Genus Melesia
21.
M. consimilis.
22. M. Scolopendrii
23.
M. Scolopendrii var. sublevis.
24. M. Darkeri.
25.
M. Dennstaedtiae.
Contrib. Arnold Arb. No. 2
Plate VII.
26
The Genus Milesia
26. M. columbiensis. 27. M. insularis.
28. M. nervisequa. 29. M. Asplenii-incisi.
Contrib. Arnold Arb. No. 2 Plate VIII
31
32
30.
M. andina.
31.
M. Histiopteridis
32.
M. tenuis.
33.
M. jezoensis.
Contrib. Arnold Arb. No. 2
Plate IX.
The Genus Milesia
34. M. vogesiaca. 35. M. dilatata.
36. M. polypodophila. 37. M. polypodophila
PUBLICATIONS PREPARED AT THE ARNOLD ARBORETUM
Of the following publications, the first can be obtained from the Arnold
Arboretum, the second and third from Houghton Mifflin Company, Bos¬
ton and New York, and the fourth from the Macmillan Company, New
York.
The Silva of North America; A Description of the Trees which
grow naturally in North America exclusive of Mexico. By Charles
Sprague Sargent. With 740 plates drawn from nature by Charles Edward
Faxon. 14 vols. f°. Boston and New York, 1891-1902. A few sets in
which some pages in vols. 4 and 14 are supplied by photostat copies.
Price $300.00
- The same. Broken sets. Number of volumes and prices on request.
- Complete sets of the 740 plates drawn for the “Silva.” Price $100.00
- Selected plates (according to number selected)
Price each $0.50 to $1.00
Trees and Shrubs. Illustrations of new or little known ligneous plants.
Prepared chiefly from material at the Arnold Arboretum and edited by
Charles Sprague Sargent. With 200 plates drawn by Charles Edward
Faxon. 2 vols. f°. Boston, 1905-1913. Price $40.00
A Manual of the Trees of North America exclusive of Mexico. By
Charles Sprague Sargent. With 783 illustrations from drawings by Charles
Edward Faxon and Mary W. Gill. Second edition reprinted with correc¬
tions. 910 pp. 8°. Boston and New York, 1926. Price $12.50
Manual of Cultivated Trees and Shrubs Hardy in North America
exclusive of Subtropical and Warmer Temperate Regions. By Alfred
Rehder. 37 + 930 pp. 8°. New York, 1927. Price $10.50
COLLECTIONS OF PHOTOGRAPHS TAKEN BY E. H. WILSON
These pictures represent trees and shrubs and different types of vegetation
found in eastern Asia, Africa, India and in Australasia ; they have been taken
by Wilson with a full knowledge of his subjects and their scientific import¬
ance. They may be obtained from the Arnold Arboretum, Jamaica Plain,
Mass.
Chinese Photographs taken 1907-1910
Series 1. 500 photographs (each 8)4 x 6)4 inches) taken 1907-1909, with
typewritten index Price $250.00
Series 2. 350 photographs (size as above, numbered 01-0350) taken 1910,
with typewritten index Price $175.00
Japanese Photographs taken 1914
A series of 600 photographs (size as above, numbered x 1-x 600) with type¬
written index Price $300.00
Photographs of the Expedition to Eastern Asia 1917-1918
A series of 700 photographs (size as above, numbered n 1-n 700) with type¬
written index Price $350.00
This series consists of the following sets : Liukiu Islands (60 photo¬
graphs) Price $30.00. — Bonin Islands (30 photographs) Price $15.00. —
Japan (50 photographs) Price $25.00. — Korea (310 photographs) Price
$155.00. — Formosa (250 photographs) Price $125.00.
Photographs of the Expedition to Australasia, Indo-Malaya and
Africa, 1920-1922
A series of 522 photographs (size as above, numbered y 1-y 522) with type¬
written index Price $261.00
This series consists of the following sets : Australasia (245 photo¬
graphs) Price $123.00. — Indo-Malaya (93 photographs) Price $47.00. —
Africa (184 photographs) Price $92.00.
Postage not included in the prices quoted above.
Selections from these photographs will be furnished at an advanced price.
PUBLICATIONS OF THE ARNOLD ARBORETUM
The following1 publications can be obtained from the Arnold Arboretum,
Jamaica Plain, Mass.
The Bradley Bibliography. A guide to the literature of woody plants,
including books and articles in the proceedings of learned societies and in
scientific and popular journals, published in all languages to the end of the
nineteenth century. Compiled under the direction of Charles Sprague Sar¬
gent by Alfred Rehder. 4°. ‘5 vols. Cambridge, 1911-1918.
Bound Price $40.00
Unbound Price $30.00
Plantae Wilsonianae. An enumeration of the woody plants collected in
western China for the Arnold Arboretum during the years 1907, 1908 and
1910 by E. H. Wilson. Edited by Charles Sprague Sargent. 8°. 3 vols.
(9 parts). Cambridge, 1911-1917.
Parts 1-4 are out of print; the remaining parts each $1.50.
The Genus Pinus. By George Russell Shaw. f°. 96 pp. 39 pi. Cam¬
bridge, 1914. Price $10.00
Catalogue of the Library of the Arnold Arboretum. Compiled under
the direction of Charles Sprague Sargent by Ethelyn Maria Tucker. f°. 2
vols. Cambridge, 1914-1917.
Vol. I. Serial publications — Authors and titles. Unbound Price $7.50
Vol. II. Subject catalogue. “ Price $7.50
The Cherries of Japan. By Ernest Henry Wilson. 8°. 68 pp. 8 pi.
Cambridge, 1916. With supplement of 3 pp. Price $3.50
The Conifers and Tax ads of Japan. By Ernest Henry Wilson. 4°.
91 pp. 59 pi. Cambridge, 1916. Price $5.00
Postage not included in the prices quoted above.
Guide to the Arnold Arboretum. 8°. 33 pp. 7 pi. 2 maps. Price $0.50
Journal of the Arnold Arboretum. A quarterly journal published by
the Arnold Arboretum. Subscriptions $4.00 per year. Price of single copies
$1.25. Back numbers on hand of Vols. VI-XII except No. 3 of Vol. X.
Bulletin of Popular Information, Series 3, illustrated. Issued during
spring and autumn, about 18 numbers per year, together with index and title-
page. Subscription $1.00 per annum. (
A limited number cf series 2, Vols. I-II (1915-16), VII (1921), X-XII
(1924-26), and of Series 3, Vol. I-V (1927-31) can be obtained at $1.00
each.
Reprints from the Journal of the Arnold Arboretum :
M
A
Northern Trees in Southern Lands. By Ernest H. Wilson. 20 pp.
1923. Price $1.00
Enumeration of the Ligneous Plants of Northern China. By Al¬
fred Rehder. — To be concluded in about 5 parts.
Part I (Ginkgoaceae to Ranunculaceae). 76 pp. 1923.
Part II (Lardizabalaceae to Rosaceae). 88 pp. 1924.
Part III (Leguminosae to Sapindaceae). 77 pp. 1 pi. 1926.
Price of each part $0.50 1
Corrections and Emendations of the Second Edition of Sargent’s V
Manual of the Trees of North America. By C. S. Sargent. 21 pp?<
1926. Price $0.10
The Spontaneous Flora of the Arnold Arboretum. By Ernest J.
Palmer. 57 pp. 1 pi. 1930. Price $0.50
Contributions to the Flora of the New Hebrides ; Plants Collected 1
by S. F. Kajewski in 1928 and 1929. By A. Guillaumin. 118 pp. 5 figs, r
1 pi. 1931-1932. Price $1.00
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